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Full text of "British Antarctic Expedition, 1907-9, under the command of Sir E.H. Shackleton, c.v.o. Reports on the scientific investigations .."


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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. ON T 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- <CAPE ROYDS* 

BY JAMES MURRAY 



Ix making the biological collections, almost every member of our small party lent 
more or less assistance. Though busy with other occupations, all were ready to bring 
home to the biologist anything strange which they noticed. Armytage found a sea- 
urchin during a walk on the sea-ice before we had made a landing, and while landing he 
picked up the first scrap of sea-weed. Mr. Shackleton brought in some moss and lichen 
soon after the Nimrod departed. Wild got specimens of the lake vegetation. Adams 
found a starfish on the beach, and the others in like manner helped when opportunity 
offered. 

All these indications that there was life in the district, coming as they did before 
the biological work was properly started, were encouraging for the future. 

When the dredging operations began there were many willing helpers. Mr. 
Shackleton, with David, Mawson, and Priestley, were always interested in the dredg- 
ing and ready to help, not only at hauling the dredge, but at the more disagreeable 
labour of conveying the collections to the hut. It is no disparagement to others to 
acknowledge the share which Priestley took in the biological work. Without him the 
greater part of the collections would not have been made. When the biologist was 
debarred by some trifling but mastering indisposition from active participation in the 
dredging during the midwinter weeks, Priestley kept the dredging-holes open, no small 
labour with the temperature sometimes as low as from minus 30 to minus 40 Fahr. 
With the assistance of Mr. Shackleton, David, and others, he kept up the dredging 
and brought home the proceeds to be examined. 

Throughout the entire season practically the whole of the arduous labour of digging 
holes in the sea-ice and of sinking shafts in the lakes fell upon Priestley, and he did the 
active hauling of the dredge as well. It does not lessen our indebtedness to him to 
tell that he enjoyed his self-imposed task, and his voice might be heard issuing in 
light-hearted song from some deep shaft in lake or sea. 

The Field of Operations. This was extremely limited. The great majority of the 
collections were made within a radius of a few miles from our base camp at Cape 

* Cape Royds is the westernmost point of Ross Island, South Victoria Land, and is situated in about 
latitude 77 32' S., longitude 1G6 12' E. 

BRIT. ANTARCT. EXPED. 1907-9. VOL. I. PART 1, ISSUED APRIL 1910 A 

13 
M749992 



2 JAMES MURRAY 

Koyds. Those who went on the long sledging journeys brought back specimens from 
more distant points. Mr. Shackleton brought lake vegetation and Joyce brought moss 
from Hut Point (the Discovery's winter quarters, twenty miles south of Cape Royds) ; 
Priestley brought rotifers, mosses, lichens, and some marine organisms from the 
neighbourhood of the Ferrar Glacier and the Stranded Moraines, when he visited the 
west with Armytage's party ; and Brocklehurst on the same journey obtained some 
lichens at an elevation of about 4000 feet at New Harbour Heights ; David brought 
moss and lichen from near Cape Irizar, the most distant point from our camp at which 
any biological specimens were collected. So far as known, the species were the same 
in all the localities. 

The promontory of Cape Boyds, round which most of the collecting was done, is a 
hilly tract of triangular form, separated from the main mass of Mount Erebus by a 
valley in which there is a series of small lakes, and terminating at its southernmost 
point in a bluff rising vertically from the sea to a considerable height. This culminat- 
ing-point of the Cape was familiarly known as Flagstaff Point, from a pole which we 
erected there for the purpose of signalling to the ship. In the hollow between it and 
the hut was Pony Lake, and between the lake and the shore was the rookery of 
Adelie Penguins. 

The triangular area is just about a mile in length and half a mile in greatest breadth. 
It includes many little sharp rocky peaks, composed of kenyte, with ridges of the same 
rock diverging from the peaks. The valleys are filled with a gravelly debris resulting 
from the decomposition of the kenyte, and contain many little lakes or ponds. Con- 
siderable stretches of morainic material occur. The highest point of land is no more 
than 300 feet above the sea. The greater part of the shore-line consists of low cliffs 
with a few small patches of sandy beach. The most extensive of these beaches, known 
as Black Sand Beach, is about a mile to the north of the hut. 

On the shore there is no vestige of marine life, animal or vegetable, such as is 
found in the littoral zone of other coasts. The beaches are formed of a coarse, hard, 
black sand, with boulders of kenyte and other rocks. The presence of an ice-foot 
throughout the greater part of the year, and the grinding of ice along the coast when 
there is open sea, must destroy any living things which attempt to establish them- 
selves. The zone thus kept devoid of life is of no great depth. Standing on the 
edge of the ice-foot at Black Sandy Beach, when the Sound was open, various living 
things could be seen at a depth of from one to two fathoms. Starfish were commonest 
in this situation, but a living Pecten Colbecki was got in equally shallow water at 
Back-door Bay. 

The larger lakes were given distinguishing names. Pony Lake, close beside the 
hut, formed the exercising ground for the ponies during the long night. A short 
distance to the north was Green Lake, named from the colour of its ice. A mile north 
of the hut and close to the shore was Coast Lake, remarkable for its level smooth 
ice, which would have served for skating and curling. Close by was Clear Lake, named 



ON COLLECTING AT CAPE ROYDS 



FIG. 1. 




M9 MURDO 
SOUND 



CAPE ROYDS 



SKETCH-MAP OF THE IMMEDIATE NEIGHBOURHOOD OF CAPE ROYDS 
Showing the positions of the lakes, &c. 

Copied from Map lent by R. E. PRIESTLEY. 



.it>-(> 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" 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 



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. 7a7e) 

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 
yy T 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 







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. y o ] j pj a t e Xlll 

MURRAY: ANTARCTIC ROTIFERA. 




16. b. 



14, PLEUROTROCHA, sp 15, FLOSCULARIA, S P . 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 
60 ime 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 7730' and 78S.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. 2a2f) 

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<* 
hatched out, and were exhibited at the Quekett Microscopical Club by Mr. > 
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 





a 




? rc 






H 














CQ 

Q 






B 


o 


^ Q 


o 


^ 











43 


^ 


w 








^ 





-^ , 


^ 'f : . 


H4 


o 








o 


u 


00 


3 






J 


a 


"^ * 


<J Q 


a 


o 




1 


^ 


*"J 


2 


3 


J 


d 






C3 






tf 








u 


H 


II 


H 


CQ 






3 

o 




CQ 

1 


is 


o 2 

3 

&S 


o 


o 




s 


o 
a 

s 


1 


3 

ftj 


K 

s 
< 




< 


2 
g 


1 t 

o 

z 


H 
o 

H 




^ 


O 


M 2 


K 


2 


55 








ty^ 


tn 


D 













o " 






cc 














I 




Echiniscns arctnmys (?) (see p. 1 2C) 








X 








X 


X 


X 


X 








X 


X 


X 


K. wendti . 








X 











X 




















X 


E. meridionalis . 








X 














































Milnesiwn tardigradum 








X 





X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Macrobiotus furciyer 








X 


X 


X 




















X 








3[. echinoyenitits 








X 








X 


X 


X 


X 


X 





X 


X 


X 


^f. meridionalis . 


X 






















































M. polaris . 


X 






















































M. oberhciiiseri . 


X 








X 





X 


X 


X 


X 


X 


X 





X 


X 


M. arcticws 


X 





. 











X 





X 


X 





X 





X 


J/. asper . 








X 





X 





























M. iiuirrayi 








X 





X 





























M. aitlarcticiis . 





X 














X 




















. 


DipJutscon cluleiiense 








X 











X 


X 


. 


X 


X 


X 


. 


X 


1). alpinum 


X 





X 











X 








X 





X 





X 


D. scoticum 







X 


X 





X 


X 








X 


X 


X 


X 


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 




, 








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 




Si 

Ed 


<y 

u 


i 


O 

*H 


1 




s 

K 


3 


s 

< 





i 






^ 


EC 


02 


S 


! 


i 








fe 


02 



































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 





!</ 'ntfmfrt'nrHji 
































X 
























M. inlermedins 


X 


X 








X 


X 


X 


X 


X 


X 


X 


X 


. 


M. dispar 





X 








X 





X 











X 


X 





M. arcticus 


. - 


X 





, 


X 





X 





X 


X 





X 


X 


M. nodosus 


X 








X 





X 








X 














M. sattleri 





X 








X 


X 


X 


X 





X 











M. papillifer 





X 


. 


. . 


X 





X 




















M. ornatus 





X 














X 


__ 











X 





M. annulatus 





X 














X 














X 





Diphascon chilenens 





X 


X 





X 





X 


X 





X 


X 


X 


X 


D. alpinum 





X 














X 








X 


X 


X 


X 


D. scoticum , 








X 





X 


X 


X 








X 


X 


X 


X 



TARDIGRADA: NEW ZEALAND 121 

The foregoing table shows the distribution of the New Zealand species over the 
world. For this purpose the surface of the globe is divided into nine great areas, 
being one for each of the continents, for the Arctic and Antarctic regions, and for 
Australasia. The islands adjacent to the continents are included with them, New 
Zealand is included with Australia, but Oceanic Islands have a column to themselves. 
The group of the " Oceanic Islands " is unsatisfactory, as the only character which 
they have in common is their isolation. The group is formed to gather up some odds 
and ends of Tardigrade records from all over the world. 

For the local distribution the Australasian column is subdivided into five sub- 
ordinate columns. 

In compiling the table I have used the published papers of Richters, Plate, and 
others, and for some regions (S. America, Africa) I have used unpublished notes 
of my own. Many records may have been overlooked, but in any case the whole 
aspect of the facts is liable to be changed at any moment by fresh work in any of 
the regions. 

As our knowledge stands, it appears that of the 21 identified species 16 are also 
found in Europe, 13 in Australia and the Arctic Region, 11 in N. America, 9 
in S. America, 10 in the Oceanic Islands, 8 in Africa, 7 in the Antarctic and 
Asia. 

These proportions indicate probably rather the amount of work done in the 
different regions than actual facts in distribution. This is especially likely to be 
true in the case of Europe, where water-bears were first studied, and most of the 
species discovered. 

Nevertheless, for what the facts are worth, the table shows that temperate New 
Zealand has a closer correspondence with temperate Europe than with any other 
country. In has as much affinity with the Arctic Region as with its nearest neigh- 
bour Australia. It has least in common with the adjacent Antarctic (which doubtless 
means that the Antarctic is poor in species) and with Asia (which is very little known). 

All the species which are common to New Zealand and the Arctic are also found 
in Europe. Of the New Zealand species which are also in Australia and in the 
Arctic, there are 10 species which are common to these two regions. 

The range of the various New Zealand species works out as follows. Milnesium 
and M. echinogenitus are in all 9 columns, M. hufelandii and M. intermedivs in 
8, D. chilenense and D. scoticum in 7, E. mutabilis and M. arcticus in 6, 
M. harmsworthi, M. sattleri, and D. alpinum in 5. The others must be considered 
local. 

E. novcezeelandice is known only in the three localities, New Zealand, Australia, 
and Hawaii, all widely separated. M. furciger is a southern species, occurring in 
New Zealand, S. America, and the Antarctic. Several others have no continuity 
in their range, as at present known. M. nodosus is in New Zealand, Fiji, and 
Africa; E. gladiator in New Zealand, Europe, and N. America; M. papillifer in 

BRIT. ANTARCT. EXPED. 1907~9. VOL. I. R 



J. MURRAY 

New Zealand, Australia, and Europe ; M. ornatus and M. annulatus in New Zealand, 
Europe, and the Arctic. Two species only, E. velaminis and M. montanus, are not 
known except in New Zealand. 

Any considerations which might be offered as to the possible origin and history 
of the New Zealand Tardigrade fauna will be reserved for the paper on Geographical 
Distribution, in a later number of this publication. 



AUSTRALIAN TARDIGRADA 

On collecting in Australia,. On the return of the Nimrod from the Antarctic 
in the spring of 1909 nearly a month was spent in Australia in the collection of 
various fresh-water microscopic animals, chiefly Rotifera and Tardigrada. 

In this work I was greatly assisted by some good friends in Sydney, Mr. E. G. 
Goddard, Mr. S. Johnston and Mr. T. Whitelegge, who guided me on excursions 
to various likely collecting-grounds in the neighbourhood. Without this friendly 
assistance the time might have been to a large extent wasted, as in a country so 
generally parched us "Australia seems to be, mosses are not so much in evidence as 
in moister countries like New Zealand. 

Only two of the Australian States were visited, New South Wales and Queens- 
land. The immediate neighbourhood of Sydney was not very productive. There is 
usually very little moss in the Eucalyptus bush, and when some is found it often 
yields few microscopic animals, or none. Some water-bears were got in moss from 
the National Park, near Sydney, and in the ponds in the Botanic Gardens some of 
the purely aquatic species were collected. 

It was only in the mountain ranges, where a moister climate prevails, and mosses 
are more abundant, that many Tardigrada were obtained. Two mountainous 
regions in New South Wales were visited the Australian Alps near the Victorian 
border, and the Blue Mountains near Sydney. 

On the journey to the Alps, the last stage of which was by coach, although the 
road ran at a considerable elevation, and the air was cool, the country still seemed 
arid. The hard baked earth showed everywhere among the gum-trees, and no moss 
was visible. At the guest-house of " the Creel," altitude about 3000 feet, on the 
Snowy River, there was scarcely any moss except on the banks of the river. This 
moss was as unproductive as that gathered near Sydney, probably because it is dried 
for too long at a time. 

With Dr. Mackay an ascent was made to the Hospice, a government guest-house 
in course of erection on the road to Mount Kosciusko, at an altitude of about 
5000 feet. As we ascended from 3000 to 5000 feet the climatic changes which we 
passed through were surprisingly great for such a small difference of level. The first 
part of the way, from 3000 to 4000 feet, was through bush, which was dry and 
barren, the pasturage for the few sheep which we saw consisting of little isolated 
tufts of grass, separated by yards of bare earth. 

123 



J. MURRAY 

The bush was cheerful with the whistling and chattering of parrots of many 
sorts and sizes, and of crows and magpies. Sometimes an isolated gum-tree would 
be seen loaded with white cockatoos, looking at a distance like blossom. As we 
approached the 5000 feet level the signs of a moister climate gradually increased. 
Mosses appeared, and became increasingly plentiful. As we topped the rise, at some- 
what over 5000 feet, we came out on a stretch of moorland, apparently covered with 
heather and familiar heath-plants and mosses. Dr. Mackay ran down and gathered 
some of the seeming heather and other plants. They were all strange and foreign, 
as was only to be expected. The purple-tipped heather was not a plant in bloom 
at all, but was something like Vaccinium with the young leaves purplish brown. 
The peat-mosses (Sphagnum) were genuine and proved that we had reached a 
temperate clime, where a rich harvest of water-bears and other animals was to be 
looked for, and this anticipation was not disappointed. 

Dr. Mackay went on and ascended Mount Kosciusko itself, but as the journey 
was made on horseback there was no chance to gather moss on the highest point of 
land on the Australian Continent. A great deal of moss was collected on lesser 
peaks, between 5000 and 6000 feet in height. The mosses afterwards yielded many 
water-bears, including two interesting species which are new to science (Echiniscus 
jmlcher and Macrobiotics aculeatus). 

On the trip to the Blue Mountains Mr. Goddard kindly accompanied me as 
guide. It was a hurried visit, as only two days could be given up to it. At 
Katoomba, where we stayed, at an elevation of something like 3000 feet, the air 
was invigorating, but the sun was hot, the earth was scorched, and no moss was to 
be seen. But Mr. Goddard had good reason for selecting Katoomba. It lies close- 
to the edge of the wonderful sunken valley, between 1000 and 2000 feet in depth, 
bounded by vertical precipices. Little gullies, leading to the valley bottom, have 
been utilised for the making of stairways, by which access is now easy. In these 
shady gullies there are trickles of water and plenty of moss, including even 
Sphagnum and Laucobryum (or a plant which looks like it). 

On the day of our arrival we descended one of the stairways, some 2000 steps, 
and, after traversing a part of the bush in the valley, ascended near the Falls of 
Leura. The trees in the valley, as well as boulders and rocks, were often festooned 
with slender hanging branches of a pleurocarpous moss. Returning home, a first 
examination of the moss was made. On glancing at the field of the microscope there 
immediately met the eye a very remarkable Bdelloid rotifer (Callidina mirabilis, 
described in a subsequent number of this publication) and several other strange 
beasts were discovered on this cursory examination, including some water-bears. It 
was an index of what we were to expect from the Katoomba collections. We had 
reached our best collecting-ground in Australia. For long afterwards the moss 
continued to produce good things whenever examined, and it was a year afterwards 
that the best find of all was made, a new generic type of Tardigrade, which has been 



TARDIGRADA: AUSTRALIA 125 

named Oreella. It is essentially an Echiniscus which lacks the armature of protec- 
tive plates, and has a soft body like Macrobiotus. 

In Queensland there was only one day available for collecting. By the advice of 
Mr. Bailey, the veteran botanist, it was spent in the bush at the quiet little station 
of Eumuudi, some distance north of Brisbane. The weather was very hot, and the 
bush was very dry. The prickly wires of the Rattan, as they sawed lines in the 
face, sometimes several at one time, were rather dangerous to the eyes, and often 
required a pause of several minutes as they were carefully picked off, one by one, 
before progress could be resumed. But here and there pendent pleurocarpous mosses 
hung from the trees, and with these I filled my pockets. In some little gullies, 
where a few pools of water indicated the existence of a stream in rainy weather, 
there were tufts of various mosses. 

These mosses afterwards proved fairly productive, both of water-bears and 
rotifers. The experience with these Queensland mosses illustrates how local 
water-bears are in their distribution, and the capriciousness of the method of 
collecting from mosses. For a long time the mosses from Queensland gave very 
meagre results, and at last, a year after gathering, a tuft in no way different-looking 
from the rest, produced water-bears enough to make up quite a fair list for Queens- 
land, including two interesting new species of Echiniscus. 

Since my return to England, Mr. S. Johnston of Sydney has been good enough to 
send me freshly gathered mosses, which have afforded an opportunity of continuing 
the study of the Australian water-bears and other animals. 

The region examined in Australia extends over about 10 degrees of latitude, from 
27 S. in Queensland to 37 S. on the borders of New South Wales and Victoria. 

Previous knowledge of Australian Tardigrada. There is very little known 
of Australian water-bears. Whitelegge's "Invertebrate Fauna," 1889 (46), has no 
reference to the group. The only record I have been able to find is in 
Ilichters' " Moosfauna Australiens," 1908 (37), in which he mentions the ubiquitous 
M. hufelandii as found at Katoomba, in the Blue Mountains. 

LIST OF THE SPECIES COLLECTED 

Echiniscus mutabilis, Murray. Milnesium tardiyradwm, Doyere. 

E. nwcezeelandice, Richters. Maerobiofus hufelandii, Schultze. 

E. pulcher, sp. n. M. echinogenitus, Richters. 

E. arctomys, Ehrenberg. M. areolatus, Murray. 

E. kerguelensis, Ricbters. M. harmsworthi, Murray. 

E. tessellatus, sp. n. M. hufelandioides, sp. n. 

E. intermedius, sp. n. M. occidentalis, Murray. 

E. spiniyer, Richters. M. intermedius, Plate. 

E. duboisi, Richters. M. crassidens, Murray. 

E. blumi, Richters. M. acideatus, sp. n. 

E. oihonme, Richters. M. dispar, Murray. 

Oreella inollis, gen. n., sp. n. M. arclicus, Murray. 



126 J. MURRAY 

M. sattieri, Eichters. M. virgatus, Murray. 

M. pupillifer, Murray. Diphascon chilcnense, Plate. 

M. rubens, Murray. D. scoticum, Murray. 

M. augusti, Murray. 

Seven species not identified (5 Echiniscus, 2 Macroliotus). 

NOTES ON THE SPECIES 

Genus Echiniscus, Schultze (43) 

A: SPECIES HAVING SEGMENTS V. AND VI. DISTINCT 

Echiniscus mutabilis, Murray (12) 

Common in all the localities visited, both in New South Wales and Queensland. 

It is desirable that a correction should be made in this place in regard to all 
records which I have made heretofore of E. arctomys in various countries. All 
these records must be taken as referring to E. mutabilis. The error arose from a 
misunderstanding of E. arctomys, Ehr. That species was supposed by Richters to 
have segments V. a,nd VI. separate, but V. was described as being a half-ring, not 
a pair, and there were no spines on the inner claws. E. mutabilis has V. paired, 
and there are small barbs, easily overlooked, on the inner claws. The differences 
are slight, as the separation of the third pair (segment V.) of plates in E. mutabilis 
is very obscure. If it is conceded that these differences are not of specific value, 
then Richters' records of E. arctomys may also refer to E. mutabilis, Murray.* 

The true E. arctomys, Ehr. [" Mikrogeologie," 1854 (4)], is quite a different animal. 
It has segments V. and VI. united (see p. 128). 

If it is sought to identify E. mutabilis with any of Ehrenberg's species it must be 
with E. suillus. The figure of that species is only a profile, and therefore not very 
satisfactory for identification. It is not clear if the last line crossing the trunk is 
the separation of V. and VI. or only one slit of the ordinary trefoil. If it is supposed 
that V. and VI. are separate, there remains another difference. Ehrenberg figures 
on the near fourth leg one very large curved spine at the base of the claws. This 
may represent the barb of the inner claws, though if so it is incorrectly drawn. In 
any case it is different from anything in E. mutabilis. A skin of 150 M contained 
three eggs of 50 M by 42 M. 

Echiniscus novcezeelandice, Richters (37) (Plate XV. Fig. 5) 

Near Sydney and at Katoomba, in the Blue Mountains, pretty frequent. 
The figure on Plate XV. is drawn from an Australian example, in order to show 
some peculiarities which are commoner in that country. These are the forking of 

* Prof, llichters informs me in a letter that all his records under the name E. arctomys should be 
E. suilhis, Ehr. The relationship of E. 'Mutabilis to E. saillus requires further investigation. 



TARDIGRADA: AUSTRALIA 127 

the dorsal spine, and the presence of little conical processes at thep ostero-lateral 
angles of IV. Occasionally these little cones occur also at segments II. and III. 
Examples of 175 and 140 AI had eggs, which measured 56 M by 40 . 

Echiniscus pulcher, sp. n. (Plate XVIII. Fig. 34) 

Specific characters. Large, red; V. and VL^ separate ; three median plates; 
V. a narrow half-ring ; pairs obscurely divided ; lateral processes a a seta, b, c, d, 
conical knobs, e a long seta ; no fringe on fourth legs, inner claws with small barbs. 

General description. Length 300 M, seta e 100 /x. The four cirri near the 
mouth have conspicuous conical bases, the palps beside them are very large ; seta a 
measures about 75 M, and has a palp or auricle at its base. The plates may be 
regarded as eleven in number (the normal number is twelve in that section of the 
genus which has V. and VI. separate), but many of the divisions are obscure. The 
head and shoulder plates, and V. and VI., are quite distinct. The three median 
have the posterior edge distinct and rounded. The second and third median are 
divided by a median line ; their anterior edge is faintly marked, and between it and 
the pairs there is an area whioh may be regarded as belonging to either. V. is a 
very narrow band, with no sign of being a pair, and sharply separated from the 
plates before and behind. VI. is not distinctly trefoliate, but there are obscure lines 
where the cut of the trefoil would be, and those of the opposite sides appear to be 
joined. Seta e springs not from the very edge of the plate but a little way up on 
the dorsal surface. 

The animal is much flattened dorso-ventrally. The plates are very finely dotted 
with pellucid dots. The claws are slender and the small barb is very near the base. 
The legs are short. 

The larva is unusually large, measuring 175 /*. It has two claws, and differs in 
no other respect from the adult. Eggs up to the number of nine have been seen in 
the cast skin, by far the largest number I have seen in an Echiniscus. 

Habitat. Among moss from the summit of a mountain, Pretty Point, in the 
Australian Alps near Mount Kosciusko, 6000 feet, April 1909. Very abundant in 
the tufts of one kind of moss, which has not been identified. Not known elsewhere. 
The skins with eggs were common, and a number of larvse were seen. 

There are about a dozen species in the section of the genus to which E. pulcher 
belongs. They have segments V. and VI. distinct. None of these species come near 
enough to pulcher to cause any difficulty in discriminating them. Only two of them 
possess seta e (E. borealis and E. islandicus) and they differ conspicuously, having 
numerous long spines, lateral and dorsal, on the body. 

E. imberbis (38) is most like E. pulcher, but the long seta is d, and segment V. 
is a pair and bears two dorsal spines. E. pulcher is the only species known to me 
which has plate V. decidedly as a half-ring, and not paired. 



128 J. MURRAY 

B : SPECIES HAVING V. AND VI. FUSED 
fJchiniscus arctomys, Ehrenberg (3) 

On technical grounds I identify an Australian species as Ehrenberg's arctomys, 
although not quite sure that it is mature. Ehrenberg's very meagre description, 
as far as it goes, fits the Australian animal. There are nine coarsely punctate plates, 
the seta a is of moderate length, there are no other setse or spines on the body, and 
no fringe on the last legs, or barbs on the claws. 

The dots in our animal appear to be pits. The lumbar plate is strongly faceted, 
having a central panel, two lateral, and one posterior, the lateral and posterior facets 
being separated by the clefts which make the trefoil. 

The length of the animal is about 230 /x, exclusive of the fourth legs ; seta a 
measures about 50 n, and the claws 12 to 15 M. 

Habitat. Katoomba, Blue Mountains ; fairly plentiful. 

As already pointed out (pp. 110 and 126), all my previous records under the 
name of E. arctomys are erroneous, and really refer to a form of E. mutabilis. This 
is the first record I have made of the true arctomys, agreeing with Ehrenberg's 
description and figure. 

According to the records E. arctomys is one of the most cosmopolitan of 
Tardigrada, but it is doubtful whether many of them refer to the true arctomys 
of Ehrenberg. 

Echiniscus kerguelensis, Richters (31) (Plate XVI. Fig. 13) 

Professor Richters, unfortunately, gives no figure of this species. From his 
description it appears to be distinguished from E. arctomys by the presence of a 
fringe on the fourth leg, and by the weaker granulation. 

As in the absence of a figure of the species the identity is not quite certain, I 
here figure and describe fully the Australian form. 

Description. Total length, exclusive of legs, 225 M, seta a about 80 M long 
(which is longer than that of the Australian arctomys, but shorter than that of 
tvendti). Plates nine, coarsely punctate with apparent perforations. The auricles at 
the bases of setse a are prominent and somewhat elongate. The mouth cirri and 
palps are normal. The plates of the pairs are each divided into two parts by a line 
which cuts off a narrow zone parallel with the anterior border. There are two 
median plates. The lumbar plate is trefoliate and is not faceted. The fourth legs 
are fairly long, and each has a blunt palp near its base. The fringe consists of 
narrow sharp spines. The inner claws have decurved barbs. The colour is red. 

Habitat. The Australian Alps of New South Wales, near the Victorian border, 
altitude 5000 to 6000 feet. 

The occurrence of this Kerguelen species in Australia, one of the nearest masses 



TARDIGRADA: AUSTRALIA 129 

of continental land, is not at all surprising, and at the high elevation at which 
it lived the climatic conditions might approximate nearly to those of the bleak 
southern island. 

Echiniscus tessellatus, sp. n. (Plate XVI. Fig. 15) 

Specific characters. Small, yellow, plates nine, Y. and VI. joined, coarsely papil- 
lose. Seta a at base of head, very long. Shoulder plate divided into ten papillose facets, 
separated by plain bands ; lumbar plate divided into six similar facets ; the paired 
plates divided into three portions each by lines merely ; two pairs, lumbar plate 
trefoliate. The lateral- ventral margins of the principal plates with a non-granular 
band, which in empty skins forms a glassy clear zone round the whole animal like a 
nimbus. Last leg with a spiny fringe, claws without barbs. 

General description. Length 200 M, seta a 150 /. The form is short and broad, 
relatively broader in the empty skin. The plain lateral marginal band of the 
shoulder, paired, and lumbar plates appears to be free from the body, forming a 
kind of projecting flange, which can sometimes be seen in the living animal, but is 
more pronounced in the empty skin. The papillae are large hemispherical knobs, the 
coarsest known in an Echiniscus. The subdivisions of the shoulder and lumbar 
plates are not to be regarded as distinct plates ; they are panels ornamenting the 
surface. The lumbar plate is faceted, having four surfaces, which meet at an angle 
a central one, two lateral, and one posterior. 

The papillae are seen on the posterior margins of most of the plates, showing that 
the skin is there bent over. 

The two-clawed larva is known, and skins with one or two eggs have been seen. 

Many species of Echiniscus have the plates more or less subdivided. The plates 
most commonly divided are the pairs, which are often crossed by one or two plain 
bands, separating the plates into two or three papillose portions. Sometimes the 
median plates are divided. There is no other species known which has such clear 
divisions of the shoulder and lumbar plates. This character alone distinguishes it 
from all other Echinisci. Other good characters are the hyaline margin to the plates, 
the very long head setae, and the very coarse granules. 

Habitat. Among moss from trees in the bush at Eumundi, Queensland, May 
1909. Abundant in one tuft of moss. 



Echiniscus intermedius, sp. n. (Plate XVI. Fig. 17) 

Specific characters. Small, hyaline or greyish. Mouth cirri with large conical 
bases. Seta a long ; no other processes on the body. Plates nine ; V. and VI. united ; 
two pairs ; three median ; each divided by transverse line into two portions ; lumbar 
plate not trefoliate. Plates covered with very wide shallow depressions, the margins 

BKIT. ANTAKCT. EXPED. 1907-9. VOL. I. S 



130 J. MURRAY 

of which make a regular reticulation. No fringe on the fourth leg : a blunt palp at 
the base of the fourth leg ; no barbs on the claws. 

Detailed description. Length, exclusive of fourth legs and head, about 150 /x ; 
seta a 40 /* There is no trace of the red colour so generally characteristic of Ecliinisci. 
The plates are the usual nine. The first and second median plates are each divided 
into two portions, of which the anterior is larger and has a rounded posterior border. 
The third median has a similar appearance, but the second portion is so obscurely 
separated from the lumbar plate that it may be reckoned as part of it. Under 
pressure there is no trace of division of the lumbar plate into a trefoil. This is a very 
unusual character. Though a number of species are figured without the slits, it is 
suspected that they may have been sometimes overlooked. They are often not 
visible in a dorsal view when the body is drawn together. The eyes are red. 

Habitat. Eumundi, Queensland ; two examples. 

Very similar animals in Hawaii and Canada show an interesting series of changes 
in the markings of the plates. In the type the reticulation is relatively large, the 
hexagons measuring 5 or 6 /* in diameter ; in the Hawaiian variety they are only 
2 or 3 M in diameter ; in the Canadian form no depressions or reticulation can be 
detected, but the plates are marked by extremely fine pellucid dots. These differences 
are of some importance, and probably indicate that the three forms have been long 
isolated. 

In Queensland neither eggs nor larva? were found. It would have been impossible 
to decide whether the specimens were mature, and the very small size would have led 
to the supposition that they were young. The Canadian and Hawaiian examples 
supplied what was lacking in our knowledge. In Canada the species was plentiful. 
Though no eggs were found, there were a number of larvre. These have a distinctive 
form, especially of the head and the lumbar plates, which are quite different from 
those of other species, but more easily understood from the figure than from 
descriptions (Plate XX. Fig. 52l>). In Honolulu the eggs were found. Skins with 
eggs were even smaller than the Australian examples, so it may be supposed that 
these were full-grown. 

The peculiar faceting of the lumbar plate, so well marked in Canada and Hawaii, 
was not observed in Australia, but the two specimens studied were not in favourable 
condition for observing such characters. 

The species may be easily distinguished from all others yet known by the small 
size, large seta a, divided median plates, lack of red colour, and of trefoil. 

The name intermedius does not indicate that the animal is supposed to be truly 
intermediate between the two great divisions of the genus (those having V. and VI. 
fused, and those having them separate). It shares some of the characters of both, 
without being genetically intermediate. 

E. intermedius belongs to a small group of species which have no dorsal or lateral 
processes after the seta a at the base of the head. There are about a dozen forms in 



TARDIGRADA: AUSTRALIA 131 

the group. All of them differ conspicuously and need not be compared in detail. It 
is enough to say that no one of them has the median plates transversely divided, 
except the variety exarmatus of E. gladiator. That has large barbs on the inner 
claws of the fourth leg, the paired plates gape behind, and there are other points of 
difference. 

Echiniscus spiniger, Richters (28) (Plate XVII. Fig. 24) 

Habitat. Among moss from the bush at Eumundi, Queensland. 

This Australian animal is identified with Richters' species, despite some small 
differences, which do not appear to be very important. Richters' type has the four 
lateral spines, b, c, d, and e, nearly equal, and from 30 to 36 p. in length ; the dorsal 
process over c is a seta of 51 M, and that over d a curved spine of 48 M. 

The Australian form has the spine b shorter, of 8 to 10 /j. in length ;. c, d, and e of 
about 30 M ; the dorsal spines over c and d are short, of only 10 to 15 /. The dorsal 
processes vary greatly in size, and may be absent. Many species vary in the same 
respect. 

The teeth or spines of the fringe on the fourth legs are obtuse. The barbs of the 
inner claws of the fourth legs are very strong, and are placed farther from the base 
than in most species, resembling those of E. gladiator. The plates are marked with 
very fine pellucid dots. Richters says nothing as to the granulation. 

Unfortunately Richters gives no figure of his species, but his description is such 
that the positions of the spines and setse can be definitely known, and a diagram of 
the animal can be constructed from it. 

Echiniscus duboisi, Richters (25) (Plate XVII. Figs. 19-20) 

Two varieties of this species occurred in Australia, neither of them agreeing 
closely with the type. That has the four lateral spines, b, c, d, and e, short, nearly 
equal, and finely spinulose ; the dorsal spines smooth. 

Description of the Australian varieties. Variety 1 (Fig. 19). Size moderate, 
length about 250 M, exclusive of the legs. Seta a is 80 ^ in length, and the " auricle " 
at its base is elongate (about 15 n in length). The four lateral spines are of different 
lengths b is 25/u long, c 46 n, d 50 M, and e 40 M, but they vary considerably in different 
individuals. The dorsal spine over c is very short (12 M), and is smooth. It is inserted 
nearer the median line of the body than the postero -dorsal angle of the plate. The 
spine over d is 50 M long, flat, and serrate on both margins. 

There are two median plates, and a dotted band behind the second pair, which 
may be reckoned as a plate or not. The dots on the plates, which Ricliters describes 
as coarse granules, seem in our specimens to be pits or perforations, irregular in size, 
and some very large. Each plate of the pairs is divided into two parts by a broad 
plain baud, ou which there are no dots. The spines of the fringe on the fourth leg 



J. MURRAY 

are narrow, acute, curved, and are separated at their bases. There are no outer or 
inner barbs on the claws. 

Variety 2. Smaller, length 190 M . Seta a not seen (only dead skins were 
examined). The three lateral spines (b is absent) are nearly equal, of 30 to 40 M. 
They are usually strongly curved, and they bear few (three or four) large spinules, 
which are also curved. The dorsal spines over c and d are of similar length to the 
lateral spines. They are broad and flat, and are irregularly serrate or merely erose 
on the margins. The dots on the plates are like those of variety 1. There is no 
note of the crossing of the paired plates by a plain band. Fringe and claws were 
not seen. 

The middle lobe of the trefoil of the lumbar plate seems marked off from the 
anterior portion by a narrow band, interrupting the granules. This may be merely 
optical, as it often happens, when this part of the plate forms a panel set at an angle 
to the rest of the plate, that the thickness of the plate is seen in optical section, 
marked by two parallel lines. 

Habitat. Variety 1 common in the Blue Mountains and in Queensland ; variety 
2 Queensland. 

The Australian varieties differ from the type in having the dorsal processes also 
serrate. Variety 1 is nearer the type ; variety 2 lacks spine b. 

Echiniscus blumi, Eichters (27) (Plate XVI. Fig. 14) 

Habitat. Australian Alps, several examples. 

Agreeing in the main with Eichters' figure, there are some small differences of no 
great importance. I saw no third median plate, but that may often be hidden by 
:he pair in front. The dots, which appear to be granules, are smaller and closer. 
Che lumbar plate is not divided right across, as in Eichters' figure, but simply divided 
by two slits into a trefoil. In this respect I believe Professor Eichters now agrees 
with my interpretation. 

Some of the setae in the example figured have little curved branches, which I 
have seen in no other species except an undescribed Antarctic one. The straight 
spines on the outer claws of the fourth legs vary from one to three in number, as I 
have previously observed in E. granulatus. The largest example measured 400 M in 
length, exclusive of the fourth legs. 

Echiniscus oihonnce, Eichters ? (27) (Plate XVII. Fig. 21) 

There are several points of difference from Eichters' type which render this 
identification more than doubtful. The lateral process c is reduced to a long spine : 
the dorsal process over d is elongated into a sharp spine of moderate length ; the 
spicules at the bases of b, c, d, and e, were not observed. In the aggregate these 
characters are of some weight, but the most important one (the lack of spicules) is 



TARDIGRADA: AUSTRALIA 133 

very difficult to be sure about, and I do not feel justified in separating a new species 
on a mere difference in the relative proportions of two processes. 

In a subsequent chapter in this paper, on Canadian Tardigrada (p. 163) another 
form is recorded doubtfully as E. oihonnce. Its divergence from the type has taken 
quite another direction. On any estimate of specific values these two extreme forms 
(the Canadian and the Australian) cannot be regarded as one and the same species, 
but we must learn more about the extent and the^ limits of variation among Ecliinisci 
before we shall know what to make of them. 

Description. Length 250 ,, exclusive of legs; seta a 130 M, & 36 M, c 50 M, 
d 60 M, e 150 M. Mouth cirri long and palps large. Dorsal processes, over c a seta of 
120 M, over d one of 30 n. Plates nine, two pairs, two median. The dots are of moderate 
size, and appear to be pits. They are uniform in size and regularly spaced. The 
lumbar plate is trefoliate and divided into four facets, of which the posterior one 
shows obscure subdivision into two. The spines of the fringe are triangular and acute. 
The claws are large (actual measurement not noted) and the inner ones have de- 
curved spines (barbs) a little way above the base. 

Habitat. Among moss, Australian Alps near the southern border of New South 
Wales, altitude 5000 to 6000 feet. 



Echiniscus, sp. ? (Plate XVI. Fig. 18) 

A large red animal (the figure is drawn from a small individual) with no dorsal 
processes. Plates nine, V. and VI. united, two pairs, two median. Lateral processes 
four setse, a, c, d, and e, each 100 M to 150 M or even more in length. Auricle at 
base of seta a exceptionally long (about 15 M), palp near mouth very large. The 
markings on the plates are larger and smaller dots, which look like perforations. 
Each of the paired plates is divided into two dotted areas, separated by a plain 
undotted band. The fourth legs have a fringe of sharp spines, and the inner claws 
have small decurved barbs. 

The texture of the plates, and the interruption of the paired plates by a plain 
band, are exactly as in E. duboisi (Plate XVII. Fig. 19) and E. spinulosus (Plate 
XIX. Fig. 38). These characters may be common to many species of Echiniscus, 
but species have not usually been examined with sufficient attention to the markings 
of the plates. The absence of dorsal processes is probably not a constant character. 

Habitat. Australian Alps, New South Wales, altitude 5000 to 6000 feet. 

The lateral setse are the same in number as in E. testudo and some related 
species, but they are ft, c, d, e, instead of a, b, c, e. E. velaminis has exactly the same 
seta; (Plate XV. Fig. 6), but differs in having two dorsal setoe at each side, in not 
having the paired plates divided into two dotted areas, in having the auricles small, 
and no barbs on any claws. 



J. MURRAY 

EcUniscus sp. ? (Plate XVII. Fig. 25) 

A small animal resembling that figured on Plate XVI. Fig. 18, and having the 
same number of lateral processes, but having also one dorsal seta on each side. The 
lateral setae are a, c, d, and e. They differ greatly in length. The seta a is 50 /x in 
length, c is 30 n, d 150 M (equal to the whole length of the body), e 70 /*. The dorsal 
seta is over c and measures 50 M. 

The plates are marked with obscure dots, the nature of which could not be made 
out. The fourth legs have a fringe of sharp spines, and the inner claws have 
decurved barbs of moderate size, which are farther from the base than usual. 
Colour pink. 

Habitat. Australian Alps, New South Wales, altitude 5000 to 6000 feet. 

This animal has some resemblance to E. oihonnce, but it lacks the process b, and 
it has c a spine and d a long seta, whereas oihonnce has c the seta and d the spine. 
It also lacks the lateral spicules of oihonnoe,. 

EcUniscus, sp. ? (Plate XVII. Fig. 22). 

A large animal with two very long setae on each side, in addition to seta a on the 
head. There are nine plates, V. and VI. united, two pairs and two median. The 
dots are large circles of uniform size and close together. The lateral setae are a, b, 
and c ; b and c are about 170 /x in length. Over c there is a dorsal seta also of 
170 n ; over d there is a flat spine of 25 M in length by 6 to 9 M across the base. 
An empty skin measured 270 /u in length by 140 M in breadth. 

The fringe has six to eight blunt rounded processes. There are no barbs on any 
claws. 

This animal may be compared with E. muscicola, Plate, and E. spitsbergensis, 
Scourfield. Plate's description (23) is very meagre, and takes no account of the 
fringe, the barbs of the claws, or the surface texture of the plates. The setae are 
a, b, c, and e, and over c there is a long dorsal seta. Our animal lacks seta e, 
and has an additional spine over d. 

Eichters (35) adds to Plate's description, noting the nature of the granulation, 
and that the inner claws are barbed. His description of the granulation fits our 
animal. When you focus high on the dots they appear as circles with central 
points ; when you focus deeper they become hexagons. 

Scourfield's description of E. spitsbergensis (45) is better, and he gives a figure. 
It has four seise, a, b. c, and d ; it has a flat triangular process over d ; the inner 
claws are barbed, and there is a fringe of rounded processes. 

This Australian animal stands near to both E. muscicola and E. spitsbergensis, 
from the latter of which it only differs by one pair of setfe and the lack of barbs on 
the inner claws. 

Habitat. Australian Alps, New South Wales, altitude 5000 to GOOO feet. 



TARDIGRADA: AUSTRALIA 135 

Echiniscus, sp. ? (Plate XVI. Fig. 1C) 

Small; plates nine,V. and VI. joined, two pairs, two median. Lateral processes a 
a curved seta of 50 /x ; d, a strong curved spine of 50 n. Dorsal processes over c a 
strong curved spine of 30 /*. Dots of moderate size, regularly spaced. Fringe 
dentate, with few small points. Inner claws with somewhat strong decurved barbs. 

The length is 200 ju, exclusive of the legs. - The nature of the dots is not 
apparent, whether granules or pits. The colour is red. 

Habitat. Australian Alps, New South Wales, altitude 5000 to 6000 feet. 

It is unlike any known species, but it is probably immature and might acquire 
other processes as it grows. 

Echiniscus, sp. ? larva (Plate XVII. Fig. 23) 

Description Small, length 120 p., exclusive of legs. Plates 9, V. and VI. 
united, two pairs, two median. Dots of moderate size, very regular, and appearing 
to be granules. 

Lateral processes a and e, a a seta of 36 p., e a curved spine of 18 M. Dorsal 
process a short curved spine of 8 M over d. There is a fringe of small triangular 
teeth, and the claws of the fourth legs have large decurved barbs. 

Habitat. Among moss from the bush at Eumundi, Queensland. 

It is usually impossible to assign a larva of Echiniscus to the species to which it 
belongs, although when a species occurs abundantly, without admixture of other 
species, and larvae are found with it, there is a presumption that they belong to it. 
Some larvae of species having very distinct characters (such as E. gladiator and 
E. reticidatus) can be recognised without difficulty. In this case no suggestion can 
be made as to the species. 

Oreella, gen. nov. (Plate XVIII. Fig. 26) 

Generic characters. Head with ten processes, as in Echiniscus ; body soft, not 
protected on back and sides by plates ; teeth, pharynx, and claws as in Echiniscus. 

Oreella has the soft body of a Macrobiotus, with the teeth, pharynx, claws, and 
processes on the head as in Echiniscus. Minor differences from typical Echiniscus 
may be of merely specific value, and will be noted in the description of the only 
species at present known. 

Oreella mollis, sp. n. (Plate XVIII. Fig. 26) 

Specific characters. Small, hyaline, back and sides papillose ; setae at base of 
head long, springing from the summits of large narrow conical processes, which also 
bear a little below the summit long narrow palps ; cirri near mouth with small 
conical bases ; anterior pair of processes narrow cones ; the palps reduced to small 



136 J. MURRAY 

hemispherical papillne ; no eyes ; trunk terminating posteriorly in little median 
process ; legs long, obscurely three-jointed ; without fringe ; claws all similar, 
without barbs ; stomach brownish. 

General description. Length, up to 230 M ; the teeth are very short, and the 
pharynx nearly as broad as long. The processes on the head and mouth correspond 
in number and position with those which characterise Echiniscus, but they differ 
in several respects. In Echiniscus each of the principal sette (or horns) of the head 
has at its base a separate triangular or oblong palp (or auricle). In Oreella this palp 
is borne near the summit of the papilla from which the seta springs. In Echiniscus 
the four processes near the mouth are all alike cirri, with minute conical bases ; in 
Oreella the anterior pair are not cirri, but narrow cones. The palps seen in dorsal 
view between the two pairs of cirri are here reduced to very small papillae. The 
body papillee are low and rounded. They might be regarded as the rudiments of 
the armour-plating, but they are not confined to definite areas, except that they do 
not extend on to the ventral surface. Each leg consists of three joints, of which the 
first and third are short, and the middle one longer. The feet are slightly webbed, 
a membrane joining the bases of the claws. Beneath the skin an obscure irregular 
areolation is seen. The stomach consists of a few large cells, which contain darker 
brown spots. No mature eggs have been seen, but in some examples there appear 
to be two very young eggs, with granular contents. 

Though in all essential features except the armour-plates very close to Echiniscus, 
there are several little points in -which it differs from all known species of that genus. 
These are, the modification of the anterior pair of processes near the mouth, the 
elevation of the auricle on the same papilla which bears the seta a at the base of the 
head, the absence of eyes, and the little median " tail" which terminates the body. 

Its movements are very slow. 

Habitat. Katoomba, in the Blue Mountains of New South Wales, altitude about 
3000 feet. It was found in moss which had been dry for about a year. When 
moistened about fifteen examples were found, of which only a few regained activity. 

Most of the genera of Tardigrada are closely related to the genus Echiniscus. 
Though no other genus may have the elaborate armour-plating which protects 
Echiniscus, all of them but Macrobiotus and Diphascon possess some of the numerous 
palps and cirri which adorn the head of Echiniscus. Several genera (Halechiniscus, 
Battilipes, Oreella) have the armature of the head almost identical with that of 
Echiniscus. Other genera have the processes more or less modified, or reduced in 
number (Microlyda [ = Lydelld], Echiniscoides, Tetrakentron, Milnesium). 

Milnesium, which departs furthest from Echiniscus, has eight palps on the head 
which may be regarded as homologous with eight of the ten processes of Echiniscus. 
In Echiniscus itself the processes may be reduced in number, though any departure 
from the normal number is very rare. 

All the exclusively marine genera, five in number, are related to Echiniscus. All 



TARDIGRADA: AUSTRALIA 137 

of these have some spines on the body which may be homologous with the lateral and 
dorsal processes, especially two at the posterior extremity which appear to be the 
most constant, and are probably homologous with seta e of Echiniscus. Even in the 
genus Macrobiotus certain species (M. dispar, M. aculeatus, and perhaps the group of 
species related to J/. tuberculatus) have dorsal processes in pairs, which may be 
homologous with those of Echiniscus. 

Oreella is nearer to Echiniscus than any of_the other genera. It has all the ten 
processes occupying the same positions, and only slightly modified in detail. The 
teeth, pharynx, and claws are all as in Echiniscus. The only important difference is 
the lack of the armour-plates. 

GENUS Milnesium, Doyere (2) 
Milnesium tardigradum, Doyere (2) 

Common in all the districts visited, in New South Wales and Queensland. 
Milnesium is the most thoroughly cosmopolitan of all Tardigrada. 
In the Australian Alps a skin was found which contained ten eggs, each measuring 
104 n in length. 

GENUS Macrobiotus, Schultze (42) 
A : SPECIES WHICH LAY ROUGH EGGS, FREE 
Macrobiotus hufelandii, Schultze (42) 

The commonest Tardigrade here as in most other places. Found in all the 
localities visited ; eggs equally common. 

Macrobiotus echinogenitus, Richters (27) (Plate XVIII. Fig. 30) 
At Katoomba in the Blue Mountains, elevation 2000 to 3000 feet. 

Macrobiotus areolatus, Murray (19) 

At Katoomba in the Blue Mountains ; only eggs seen. This species, which was 
regarded as a variety oi'M. echniogenitus, is described as a species among the Canadian 
Tardigrada in this paper, as in Canada material was got for a study of the egg, the 
young, and the adult. It can be identified from the egg alone. 

Macrobiotus harmsworthi, Murray (19) (Plate XVIII. Fig. 31) 

In the Australian Alps, near Mount Kosciusko, altitude 5000 to 6000 feet. Not 
to be identified with certainty from the adult alone or from the egg alone. It is 
necessary to find an egg containing a well-grown young, or an adult with ripe eggs 
in the body. In this instance the young was found in the egg. 

BRIT. ANTARCT. EXPED. 1907-9. VOL. I. T 



138 J. MURRAY 

Macrobiotus hufelandioides, sp. n. (Plate XVIII. Figs. 29a-29c) 

Specific characters. Small. Claws of hufelandi type, very thick, the larger one 
of each pair with two strong supplementary points. Teeth slender, curved or bent. 
Gullet slender. Pharynx shortly oval, with three short rods in each row of thicken- 
ings. Egg-processes conical with expanded discoid tops. 

Detailed description. Length, 350 /" and upwards. No figures noted for larger 
examples. The teeth are slender, and taper very gradually. They are bent at a very 
obtuse angle just after emerging from the guides, and again more abruptly near 
the furca. The gullet is scarcely more than one-third of the relative width of that 
of hufelandii. The three rods in the pharynx are about twice as long as broad, 
and have rounded ends. They are of nearly equal size. The first one, though 
clearly distinguishable from the apophysis, is apparently closely attached to the end 
of the gullet. There is no comma, or a very small and obscure one. 

The stomach is brown, with darker brown patches. The fat-cells are 8 to 10 M in 
diameter. There are two dark eyes. The claws are very stout and strong ; the 
lesser one considerably shorter than the other. 

Eggs were found in the body which, when squeezed out and freed from the 
surrounding membrane, were found to have processes like those of the typical 
hufelandii egg. The processes taper from a broad base, which is surrounded by a 
circlet of dots, and are abruptly expanded at the summit into flat discs. No eggs 
were found which contained embryos, but the establishment of the relation of the egg 
to the animal was made in the more satisfactory way. If an embryo with a slender 
gullet were found in an egg, apparently of hufelandii, it might be supposed that the 
young of that species had the gullet more slender (although I believe this is not so). 
When the eggs are got in a mature adult, however, there is no such doubt. The 
species occurred in two localities in the same district, confirming the first observations. 

Habitat. Australian Alps, near the southern border of New South Wales ; at 
the Creel, altitude about 3000 feet, and on the summit of a peak near the Hospice 
on the road to Mount Kosciusko, altitude 5000 to 6000 feet. 

The combination of an egg exactly like, and claws sufficiently like, those of 
M. hufelandii, with slender teeth and gullet, distinguishes the species from all others. 
It is impossible to say whether it is really very close to M. hufelandii, or has acquired 
the characters in which it resembles it independently. All the closest relations of 
M. hufelandii have very wide gullets and thick powerful teeth. One form of M. inter- 
medius, which is not at all closely related to M. hufelandii, has similar egg-processes. 

It has been known that some forms of M. hufelandii could not be positively 
distinguished from allied species, and the egg had to be relied upon for their 
separation. It was supposed that the egg, with its disc-topped processes with their 
basal circlets of dots, was unmistakable. 



TARDIGIiADA: AUSTRALIA 189 

There must now be added this additional worry to the troubles ot the student of 
water-bears, that when he finds an egg of the hufelandi type, he must get an 
embryo in order to complete the identification. 

Macrobiotus occidentalis, Murray ? (Plate XXI. Figs. 54a-54e) 
(See Canadian Tardigrada, p. 169 in this paper) 

In the Australian Alps there occurred, at an- elevation of between 5000 and 6000 
feet, an animal closely related to that which is described under the above name in a 
later section of this paper, if it is not identical with it. 

It is a large animal of yellow colour, with dotted skin. The skin is double, with 
a clear fluid circulating between the layers. In the type this fluid contains numerous 
thin, hyaline rectangular plates, which are absent from the Australian form. There 
is a comma in the pharynx. 

The species was described among the Canadian Tardigrada, as the eggs were 
found there, and the stud)' completed. 

No eggs like the Canadian ones were found in Australia. 

An identical water-bear was got in Hawaii (see p. 155 and Plate XIX. Fig. 39.) 

The curious distribution will be discussed in the Canadian section of this paper. 

Macrobiotus intermedius, Plate (23) 

Habitat. Eumundi, Queensland. 

This species is one of a group of three, very closely related, and most readily 
distinguished by their eggs. M. intermedius has an egg with short blunt processes. 
In the typical form these are expanded from narrow bases, and are thus somewhat 
top-shaped. Another form of egg is like a miniature of that of M. hufelandii. The 
processes taper from the base and are surmounted by disc-shaped or saucer-shaped 
expansions. 

The egg of M. crassidens has very long slender processes, that of M. aculeatus 
has somewhat similar, but thinner and more scattered, processes. That animal is 
distinguished by the dorsal processes. 

Sometimes the processes of the variety having eggs of the hufelandi type are 
irregularly furcate at the tips, recalling those of M. furciger and M. orcadensis. 

Macrobiotus crassidens, Murray (20). 

At Katoomba in the Blue Mountains, New South Wales, altitude 2000 to 
3000 feet. 

Macrobiotus aculeatus, sp. n. (Plate XVIII. Figs. 27a-27e) 

Specific characters. Size moderate ; old examples densely pigmented ; skin 
bearing 2-6 soft white conical dorsal processes in pairs on the segments over the 



140 J. MURRAY 

second, third, and fourth pairs of legs; teeth slender, curved, gullet narrow, expanded 
in the pharynx ; pharynx nearly round, containing apophysis, three very broad 
nuts, and an obscure "comma" in each row. Claws of the hufelandi type, 
slender, unequal in each pair, and united for half the length of the longer one. 
Egg small, covered with very slender undulate or curved processes, which spring 
from small conical bases, and are separated by interspaces greater than their own 
diameter. 

General description. Considered as one of the smaller Macrobioti, though 
individuals occasionally attain to a very considerable size. Length of an ordinary 
example about 300 /u. Only the very large individuals are strongly pigmented ; 
those of moderate size are generally nearly colourless except in the stomach. The 
pigment is not in bands. There are usually two small dark eyes, but they are 
sometimes absent. The nuts in the pharynx are often broader than long. The first 
appears to be closely united to the gullet, though distinct from the apophysis. The 
dorsal processes can be seen in the young squeezed out of the ripe egg. 

M. aculeatus is very closely related to M. crassidens, Murray (20), and to 
M. intermedius, Plate (23). The spines distinguish it from these species. It has no 
close affinity with M. papillifer, Murray (12), and M. sattleri, Eichters (26), which 
have somewhat similar spines. They belong to a different section of the genus. 
The close relationship to M. crassidens is further indicated by the similar egg. 
They are of almost identical size, but that of M. crassidens, which has been recently 
discovered in Africa, has spines with larger bases and no spaces between them. 

Habitat. Among moss ; the type with six spines in the Australian Alps near 
Mount Kosciusko, at an elevation of 5000 to 6000 feet ; a variety with only the last 
pair of spines (Fig. 27e) at Katoomba in the Blue Mountains of New South Wales. 



Macrobiotus dispar, Murray (16) 

Habitat. Ponds in the Botanic Gardens, Sydney. One of the two aquatic 
species obtained, the other being M. augusti. 

B: EGGS WITH THICK SHELLS, ENCLOSING EODS 
Macrobiotus arcticus, Murray? (19) 

Habitat. Katoomba, in the Blue Mountains. 

Without the egg the identification of M. arcticus cannot be absolutely certain, 
but the animal agreed in all other characters with this species. 

It measured 300 P. in length, the claws were of the Diphascon type, the gullet 
was slender and expanded in the pharynx, which contained two rods of nearly equal 
length and about 2|- times as long as broad, and no " comma." 



TARDIGRADA: AUSTRALIA 141 

C : SMOOTH EGGS LAID IN THE CAST SKIN 
Macrobiotus sattlcri, Richters (26) 

Habitat, Tree-moss, Eumundi, Queensland. 

Professor Richters neither describes nor figures the pharynx of this species. He 
probably found it in the " simplex " condition. The structure of the skin is very 
distinctive, and taken in conjunction with the divergent claws there would seem to be 
little difficulty in recognising the species, without considering the pharynx. There is 
reason, however, to believe that more than one form has skin like sattleri and also 
divergent claws. One of them is figured in Scottish Tardigrada (18) Plate IV. 
Fig. 2G/;. 

The Australian form has a different pharynx. There are two rods the one next 
the gullet about two and a half times as long as broad, and slightly constricted in the 
middle, making two equal lobes ; the second about twice as long as broad. There is 
no " comma." The claws are unequal, and are joined at the bases only. They diverge 
at less than a right angle. The long claw of one pair is longer than that of the other, 
and has a small supplementary point. Eyes are present. 

Macrolnotus papillijer, Murray (12) 

Habitat. Among moss from the Australian Alps, altitude 5000 feet, New South 
Wales. 

Macrobiotus rubcns, Murray (17) 

Habitat. At Katoomba, in the Blue Mountains, altitude between 2000 and 
3000 feet, abundant. 

It was not seen alive. The fat-cells still retained the red colour. The largest 
example measured 450 ju in length. No comma was seen in the pharynx, but it is often 
difficult to see in dead animals. 

The distribution of the species, so far as known, is peculiar. Discovered in India 
(17) at an elevation of 6000 feet, Professor Richters afterwards found it in Ascension 
and Sumatra (39). It now reappears in Australia, where the eggs as well as the 
adults were found. 

Macrobiotus augusti, Murray (18) 
Habitat. Ponds in the public parks, Sydney 



142 J. MURRAY 

D : EGGS UNKNOWN 

Macrobiotus virgatus, Murray ? (Plate XVIII. Fig. 28) 
See description in Canadian Tardigrada, p. 173, and figure (Plate XXI. 

Figs. 55a-55c). 

The Australian form is not fully studied, but it agrees with the type in the 
important characters of the pharynx and claws. The teeth are straight for three- 
quarters of their length, then very abruptly bent outwards. There is some individual 
variation in this respect. The pharynx possesses a minute comma, which is not noted 
for the type. There are no eyes, while the type has eyes. There are no notes as to 
the colour of the Australian examples. 

Habitat. Katoomba, in the Blue Mountains, New South Wales. 

Macrobiotus, sp. ? (Plate XVIII. Figs. 32o-32c) 

Description. Large, 450 M in length. Gullet slender ; teeth slender, abruptly bent 
towards the furca. Pharynx shortly oval, with two rods in each row, and no 
comma. The first rod is twice as long as the second, and about four times as long as 
broad. The claws are V's, or more properly U's, united at the base only. The pairs 
are unequal, and one claw of each pair is longer. The longer claws have two fine 
supplementary points. Each foot has a prominent boss near the extremity. The skin 
is finely granular. 

Habitat. Katoomba, Blue Mountains, New South Wales. 

This appears to be a very distinct specieSj but till more specimens are examined, 
and the eggs found, it is not desirable to name it. Plate figures such bosses on 
the legs of M. oberhauseri. 

Macrobiotus, sp. ? (Plate XVIII. Figs. 33a-33c) 

A small animal, resembling M. hufelandii in claws and pharynx, yet with several 
points of difference. 

Description. Very small ; the example studied measured only 120 /m in length. 
The gullet is wide, and has a very prominent rim on the end in the pharynx. The 
teeth are strongly curved. The pharynx is shortly oval, and has two rods in each 
row and no " comma." The rod next the gullet is about three times as long as broad, 
the second rod about twice as long as broad. The claws are slender, and are united 
for rather more than half their length. The two claws of each pair are slightly 
unequal. Eyes are present. 

Habitat. Eumundi, Queensland. 

Although very similar to M. hufelandii, the characters of the pharynx are 01 some 
value. It would be necessary to find the egg in order to settle the position of the 
animal. 



TARDIGRADA: AUSTRALIA 143 

Genus Diphascon, Plate (23) 
Diphascon chilenense, Plate (23) 

In the Australian Alps near the borders of New South Wales and Victoria, 
altitude 5000 to 6000 feet. 

Diphascon scoticum, Murray" (11) 

Synonym. D. crozetense, Richters (35). 

At Katoomba, in the Blue Mountains, elevation 2000 to 3000 feet, frequent. 

REMARKS ON THE AUSTRALIAN TARDIGRADE FAUNA 

Its composition. Thirty-one species have been recognised and named, but a good 
many others have been observed, though they could not be named. The list includes 
eleven species of Echiniscus, one of Milnesium, two of Diphascon, sixteen of 
Macrobiotus, and Oreella, a new genus. 

This Tardigrade fauna shows many peculiarities. It is, of course, very imperfectly 
known, but taking the facts as they stand, the following are points of interest. 
Though much less carefully worked than New Zealand, the Australian list is longer 
than the New Zealand one. 

It presents much greater peculiarity. Only about half the species are common 
or widely distributed. There is one interesting generic type, somewhat intermediate 
between Echiniscus and Macrobiotus, at present unknown elsewhere. There are six 
new species, of which five are as yet only known in Australia, and there are several 
the distribution of which is very restricted or very peculiar. 

The list is the longest one compiled for any one country during the expedition. 
In Australia we got 38 species, of which 31 were identified. Canada comes next 
with 31 species (23 identified), then New Zealand with 25 species (21 identified). 

The subjoined table shows the distribution of the Australian water-bears over 
the world, in 13 columns, 4 for the principal localities visited in Australia, and 9 for 
the rest of the world. 

The local distribution is remarkable on account of the very little indication there 
is that the species are to any extent generally distributed in Australia. No fewer than 
twenty-three out of the thirty-one species occurred in only one of the four localities 
visited. Those localities certainly offer very different climatic conditions. The 
contrast between the cool moist mountain-tops of the Alpine region at elevations of 
from 5000 to GOOO feet above the sea, and the low-lying parched bush of sub- 
tropical Queensland, is very great. The Blue Mountain region is of intermediate 
character, and it was in it that the greater number of the species, and the most 
peculiar species, were obtained. That the distribution is very local, even in any one 
district, is clearly shown by our experience in washing the moss. For a long time 



144 



J. MUMRAY 



DISTRIBUTION : LOCAL AND GENERAL 

Australia 













a 


Q 


















SH 


4 

H 





'{, 


5 


s 

J 


o 

M 





ti 




4 


o 


d 

H 




M 


J^ 


^ 


3 


3 


en 


M 


3 


CM 


^ 


o 




o 




Q 
X 




g 


1 


m 

H 


a 

H 


t i 
o 


X 




i 


o 


BD 


-rj 


o 
9 


H 

2 




02 


<} 




H 


^, 


z 




^ 


H 






"s 


M 






M 


< 


b 
9 


* 


5 


"A 


co 
























5 




























O 
















Echiniscus mutabilis 


X 


X 


X 


X 


X 


X 








X 


X 


X 


X 





E . T jiovcezeelandice .... 


X 


X 








X 


X 























E. pufoher ; 








X 
































E. arclomys '(?)*. 





X 











X 








X 


X 


X 


X 


X 


E. kerguelensis .... 








X 








X 


































X 








^_ 




















E. intermedius .... 





X 





X 





X 


X 


. . 

















E. spiniger ..... 











X 














X 














E. duboisi 





X 





X 

















X 


X 








E. blumi ..... 








X 

















X 








X 





E. oihonnce ..... 








X 











X 





X 








X 





f\ 77 77 * 




























Ureeaa tnowis ..... 




























Milnesium tardigradum . 


X 


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 


X 





M. echinogenilus .... 





X 








X 


X 


X 


X 


X 


X 


X 


X 


X 


M. areolatus ..... 





X 











X 


X 





X 


X 


X 


X 





M. harmsworthi .... 





X 


X 





X 


X 


X 





X 





X 


X 





M. hufelandioides .... 





. 


X 











. 




















jl/. occidentalis .... 








X 








X 


X 




















J/. intermedius .... 





X 








X 


X 


X 


X 


X 


X 


X 


X 


X 


J/. crassidens . . . . , 


. 


X 











X 





. 








X 








M. aculeatus ..... 





X 


X 











. 




















JJ/. dispar ..... 


X 











X 








X 


X 


X 





X 





3/. arcticus ..... 





X 








X 





X 




X 





X 


X 


X 


J/. sattle>-i ..... 


- 








X 


X 


X 


X 





X 


X 











.J/. papillifer ..... 








X 





X 











X 














jTf. rubens ..... 





X 











X 











X 


X 








.if. august! . . . 


X 























X 








X 





M. virgatiis (?) .... 





X 














X 














X 





Diphascon chileneit&e 








X 





X 





X 


X 


X 


X 





X 


X 


D. scolicum ..... 


~ 


X 


~ 


~ 


X 


X 


X 


X 


X 


~ 


~ 


X 


X 



there was very little got in the Australian moss, and then, a year after gathering 
some little tufts proved very productive. 

About a, dozen of the species are widely distributed or cosmopolitan. These occur 
in six or more of the principal columns in the table. Some are more local, and a few 
have a very restricted range. E. novcezeelandice is at present only known from 
Australia, New Zealand, and Hawaii, E. kerguelensis occurs only in Australia and 
Kerguelen. Two species, E. intermedius and M. occidentalis, are on both sides of the 
Pacific, in Australia and British Columbin, and also on islands in mid-Pacific. 
E. bin-mi is found in Australia, Europe, and the Arctic, E. oi/io/titfr in the same three 
* All the records of E. arctomys are wrong, except for Australia (see p. 120 and footnote). 



TARDIGRADA: AUSTRALIA 145 

regions, and also in Canada. M. crassidens is only known in Australia, the Pacific 
Islands, and Africa ; M.papillifer in Australia, New Zealand and Europe ; M. virgatus 
in Australia and Canada ; M. rubens in Australia, India, and some islands. 

The list of Australian Tardigrada corresponds with those of other regions in the 
following proportions: Of the 31 species, 18 are known also in Europe, 16 in the 
Oceanic Islands, 15 in the Arctic, 14 in N. America, 13 in New Zealand, 12 in 
Asia and Africa, 7 in S. America and the Arctic. 

That there is so nearly the same degree of correspondence with such diverse 
regions as New Zealand, Canada, Asia, and Africa, is an indication of how little 
proximity counts in determining relationship, and the number of rare species which 
are common to Australia and various distant countries gives some idea of the com- 
plexity of the problem of the origin of the Tardigrade fauna of any one land. 

One-sixth of all the species are peculiar to Australia. This is a large proportion, 
but a greater number of species, and a much higher proportion of them, are peculiar 
to Canada. The degree of peculiarity is much higher in Australia than in Canada, 
as the list includes one new genus, and the species are more strongly marked ones. 
It is evident that at the present time we know far too little to come to any definite 
conclusions as to the origin of the Australian Tardigrada, and their interrelationships 
with those of other countries. 



BRIT. ANTARCT. EXPBD. 1907~9. VOL. I. 



TARDIGRADA OF SOME PACIFIC ISLANDS 

FIJI : ISLAND OF VITI LEVU 

On collecting in Fiji. The collecting in Fiji was done tinder unfavourable 
conditions. The steamer stayed only for a few hours, and there was no time to get 
advice as to the most suitable localities in the neighbourhood of Suva. It was 
oppressively hot, and everything seemed burnt up by the sun. I first tried the shore 
road, skirting a mangrove swamp for some distance, but found no moss or similar 
plant. Then I made a short tour inland, passing through some banana plantations, 
and visiting the reservoir. No moss was found in all this round, although some 
patches of wood were examined. Then, returning disappointed by one of the main 
roads, I came on a little ditch and a small trickle of water connected with it. A few 
tufts of unpromising moss, silted up with gritty mud, were got alongside the ditch, 
and in some scrubby bush bordering the streamlet there were some mosses on fallen 
trees. This last was the only likely moss for the kind of animals I was seeking, and 
it was very little in quantity. The moss from the ditch was very different from the 
" dry " moss which is the happy hunting-ground of the student of " Moosbewohner " 
(moss-dwellers) ; it was likely to contain only those truly aquatic species which are 
not accustomed to being dried up periodically, and which were therefore unlikely to 
survive the long journey which had to be made before they could be examined. 
Nevertheless, when moistened and examined some months later, these unpromising 
mosses were found to harbour a considerable number of microscopic animals. These 
were chiefly Bdelloid Rotifera ; of these were about as many species as we got in 
Hawaii. There were very few Tardigrada, only two species being identified. 

The Fiji Islands are situated in the Tropics, 16 south of the Equator, and due 
north of New Zealand, which is distant about 1200 miles. The nearest part of 
Australia is about 1500 miles distant. Though so far separated from these nearest 
considerable masses of land, there are numerous intervening islands which would 
assist migration, so that some correspondence with the microfauna of Australia and 
New Zealand would not be surprising. The Tardigrada of Fiji are almost unknown. 
We identified only two species, the cosmopolitan M. areolatus (a doubtful identifica- 
tion), and M. nodosus, a species found in New Zealand. 

Previous knowledge. I can learn of no previous work on the Tardigrada of Fiji. 

147 



148 J. MURRAY 

LIST OF SPECIES 

Macrobiotus areolatus, Murray ? 
M. nodosus, Murray. 
Macrobiotus, sp. ? 

Macrobiotus areolatus, Murray ? (19) (Plate XIX. Fig. 40) 
See description of M. areolatus on p. 167 of this paper 

The identification rests on the egg only, and as the egg is by no means typical 
there is much room for doubt as to the species. The egg is considerably smaller, 
measuring only about 70 /* in diameter, exclusive of the spines. M. areolatus 
measures about 100 n without the spines. The processes also differ in form, being 
ovate and subobtuse, while those of areolatus type are conical and acuminate. They 
measure only about 12 n, while those of the type attain to 50 n in length. 

The surface of the shell, between the processes, is marked by chambers or areoltc 
like those from which the type was named, but these are more rounded and scarcely 
polygonal at all. It is on this character that the provisional identification is made. 
It is more probable that the egg belongs to a related species not yet recognised, 
perhaps to that found associated with it, and described below (figured on Plate XIX. 
Fig. 43). 

Macrobiotus nodosus, Murray (20) 

The only abundant species in the collection from Fiji. Young and old were seen, 
and skins containing the reddish brown eggs. One of these, containing three eggs, 
measured 400 n, the eggs 68 /". The largest animals were as large as African 
specimens (500 n and upwards). The claws were 30 A* in length. 

The newly hatched young of 160 ju in length were hyaline and colourless, or very 
faintly yellow. The characteristic knobs were not developed, and the papillae were 
wide but very obscure. The pharynx has the thickenings much shorter than in 
the adult. They are as broad as long, and somewhat quadrate. 

The empty skins lose the papillae, but show their position by a regular reticulation. 
Seven eggs were seen in one skin. 

The occurrence of M. nodosus in mid-Pacific is very interesting. The species is at 
present known in Africa, New Zealand, and the Macquarie Islands. 



Macrobiotus, sp. ? (Plate XIX. Figs. 43a, 

A fairly large animal, nearly 500 n in length. It is pigmented with a dull brown 
colour like that of M. Imfelandii. The teeth are stout and the gullet wide. The 
pharynx is shortly oval, and contains, besides the apophyses, three short thick rods 
in each row, and a large comma. The middle rod is a little shorter than the two 



TARDIGRADA: PACIFIC ISLANDS 149 

others. The claws are in pairs, united at the bases only. One claw of each pair is 
larger. No supplementary points were seen. There are no eye-spots. 

The pharynx is 40 /* long by 30 /u wide. Having points of affinity with several 
species (M. echinogenitus, M. areolatus, &c.), the recognition of the egg is necessary for 
its identification. The strong comma distinguishes it from areolatus, but, as Richters 
has pointed out, we are not yet sure of the constancy of the "comma." The slight 
shortening of the middle rod may show some degree of relationship to M. virgatus, a 
Canadian species (see p. 173 and Plate XXI. Fig. 55). 



HAWAII: ISLAND OF OAHU 

Collecting in Ilaivaii. At Honolulu we remained for twenty-four hours, so that 
there was a much better opportunity for collecting than we had in Fiji. As, how- 
ever, we arrived at 5 P.M. it was impossible to get clear of the town before sunset, 
and serious collecting had to be deferred till next day. The remaining daylight of 
the first evening was occupied in making a journey by street-car to the Aquarium. 
On the way some irrigation ditches were seen, and these were visited, and a net 
thrown in to collect the aquatic rotifers, with lively hopes of getting Trochosphcera, 
and who knows what else ? A few sweeps of the net filled it with a delightful green 
slime, of endless possibilities, but alas ! in drawing it out it caught on some spiny 
twig and was torn, and the contents lost. 

Next day a start was made in the cool of the morning, before sunrise. Even at 
that early hour (5 A.M.) a cafe was found open, and by the time breakfast was over 
the street-cars were running. Knowing nothing of the geography of the country, an 
electric tram was boarded, which, as luck would have it, conveyed me five miles out, 
to the foot of a valley on the east of the city. For a mile or two the road ran 
through the level valley, among farms. A very little stream ran in this valley. It 
would be only two or three feet across and a few inches deep. None of the water 
was wasted, each farm or garden using it for irrigation, and letting the overflow pass 
on to the next. No mosses were seen in this valley till the last of the houses was 
left behind, when some tufts growing on clay were got on the banks of the stream. 

The stream was now left and I began to climb up a long spur rising gradually 
towards a ridge which formed the watershed at the head of the valley. The country 
was open, with deep grass and occasional bushes. Here, at some little elevation, 
moss of a likely kind began to appear, first on stones, then on the stems of trees and 
bushes. Everything was dry and scorched by the sun. 

As I advanced the bushes became denser and rather troublesome. The moss also 
became quite abundant and my pockets were bulging with it. There were many 
kinds of it, but one was dominant. It formed large sheets of a vivid green, with 
rich autumnal browns in the withered parts. It had all the habit of a pleurocarpous 



150 J. MURRAY 

moss, but the hairy calyptra of the capsules showed it to be a relative of the Ortho- 
tricum family. 

It was my intention to go on to the top of the ridge, which seemed but a little 
way in front, and to descend into the Pali road, which I knew was only a little way 
down on the other side. Nature did not permit this. As I advanced the bushes 
became very dense and I got into a zone of ferns. The spur had become high and 
narrow, with very steep sides and scarcely walking room on the summit. 

Though there were few prickly bushes the long trailing branches of some of them 
were very tough and difficult to negotiate. The fern got so dense that it was 
difficult to part it. This innocent-looking fern appears to be a bracken, not unlike 
the British kind, which can be obstructive enough on a hill-side. It was all dry and 
withered, and the pinnules were mostly broken off, leaving only about an eighth of 
an inch of the base. These fragments of pinnules were strong and_hard, and with 
their ragged edges they caught and tore the clothing as Smilax prickles would do. 

At last, not many hundreds of yards from the ridge towards which I was making, 
the fern was so dense that I never set foot to ground at all, but walked on the 
entangled fern, a yard or more above ground. It was evident that, at the rate at 
which I was progressing, if I held on till I gained the Pali road I would miss the 
steamer, so I turned back by the way I had come. On descending from the spur I 
struck the stream at a point farther up the valley than where I had left it, and 
crossed some boggy places where there were plenty of bog Hypna and even 
Sphagnum. Had it been possible to examine these while fresh there would doubt- 
less have been interesting additions to the list of species. 

The trip was successful, judging by the quantity of moss collected, and for more 
than a year afterwards it continued to yield plenty of microscopic animals. Among 
them there were more than a dozen species of Tardigrada. 

The Sandwich Islands are situated in the Tropics, Honolulu, on the Island of 
Oahu, being 22 north of the Equator. It is the most isolated land visited. The 
nearest land mass is N. America, the nearest point being 2000 miles distant. 
Although the numberless islands of Polynesia lie to the southward, there are few 
islands near the Hawaiian group which could facilitate migration. The microfauna of 
the group may thus be expected to be of great interest. 

Previous knoidedge of Haivaiian Tardigrada. The only reference to Hawaiian 
Tardigrada with which I am acquainted is in Richters' " Moosfauna Australiens " 
(37). In moss from Oahu and Hawaii, collected by Professor Schauiiisland in 
1896-7, Richters found E. arctomys and M. hufelandii, both in Oahu, the latter 
also in Hawaii. We found the Macrobiotus again, but not E. arctomys, unless the 
record of that species refers to the same animal which I identify as E. mutabilis. 
The matter requires clearing up, 



TARDIGRADA: PACIFIC ISLANDS 151 

LIST OF SPECIES 

Echiniscus mutabilis, Murray. Macrobiotus hufelandii, Schultze. 

E. novazeelandice, Richters. M. occidentalis (?), sp. n. 

E. intermedius, sp. n. M. intermedius, Plate. 

E. viridis, sp. n. M. crassidens, Murray. 

E. spimdosus, Doyere. M, oberMuseri, Doyere. 

E. perarmatus, Murray. Diphascon scoticum, Murray. 

Milnesium tardigradum, Doyere. 

Three species not identified (1 Echiniscus, 2 Macrobiotus) 

Genus Echiniscus 
A : SEGMENTS V. AND VI. SEP ABATE 
Echiniscus mutabilis, Murray (12) 

The commonest species in Hawaii. The form of Echiniscus recorded below as 
E. novcezeelandioi has the characteristic dorsal processes so reduced in size that it 
may be questioned whether it should not rather be regarded as a form of E. mutabilis. 
At any rate it is a transitional form from the one species to the other, and indicates 
how closely related they are, although their types differ so conspicuously. 

Echiniscus novazeelandice, Richters (37) (Plate XIX.) Fig. 35 

The form of this species which occurred in Hawaii differs from the typical 
examples found in New Zealand and Australia in the extreme reduction of the dorsal 
processes on the third pair of plates. They are mere knobs or sometimes only angles 
of these plates. 

Only a few examples were seen. They measured about 240 /* in length, exclu- 
sive of the legs. The setse a were more widely spreading than usual, and were 60 n 
long. 

B: SEGMENTS V. AND VI. UNITED 

Echiniscus intermedius, Murray (Plate XIX. Figs. 36c, 36d) 
See description in Australian Tardigrada, p. 129 of this volume, also in Canadian 

Tardigrada, p. 161 

This most interesting little species occurs in Hawaii as a peculiar variety. The 
type as found in Queensland and the variety found in Canada differ so much that 
they would be considered as distinct species, were it not for the Hawaiian variety, 
which is intermediate between them. The type has the plates marked with large but 
very faint hexagons, the Canadian variety has very fine pellucid dots ; the Hawaiian 
form has markings of intermediate size, but they are depressions like those of the 



152 J. MURRAY 

type. It is a question whether all three are not distinct species. There is no other 
Echiniscus which shows such a range of variation of the surface texture. 

Description. Small, length about 190 to 220 M. Plates ten, two pairs, three 
median. All the median divided by transverse lines into two nearly equal portions. 
Both parts of the first median are dotted. In the second and third median only the 
front portion was seen to be dotted. The posterior half of the third median is only 
separated by a faint line from the lumbar plate. The plates of the pairs are each 
traversed by a transverse furrow, separating anterior and posterior convex portions. 
It is uncertain if the dots cease in the furrow. The lumbar plate is not trefoliate. It 
is faceted, having a median and two lateral panels. 

The surface of the plates is marked with a fine reticulation, regularly hexagonal. 
This is formed by the margins of contiguous shallow pits, as in E. reticulatus. These 
are much smaller than in the Australian form of the species. It must be understood 
that these pits are extremely difficult to observe. To show them by lines in the 
drawing exaggerates their apparent importance. 

The seta a, at the base of the head, is relatively long. The smallest example 
observed, 190 M in length, contained two nearly round eggs 40 p. in diameter. An 
example of 21 i contained one narrow egg of 50 n by 32 /*. 

Echiniscus viridis, sp. n. (Plate XIX. Figs. 36a, 366) 

Specific characters. Large, stout ; plates ten, three median, two pairs, V. and 
VI. joined ; colour of plates olive-green, dots darker green, regularly spaced, largest 
in the centre of the body, and diminishing to the sides ; seta a very short, springing 
from large conical papilla ; fringe of small teeth ; claws very large, inner with small 
barbs ; small spine on first leg. 

General description. Length, 250 M, exclusive of the large stout legs, claws 25 n 
long. The plates are very distinct, with clearly marked edges. Only the plates are 
coloured green ; the interior of the body is of the usual Eclviniscus red. The colour 
does not affect the bands of skin connecting the plates, but it does extend into the 
fourth leg as far as the fringe ; beyond that is clear. The dots do not appear as 
either pits or papillae, but simply as darker patches. They are very regularly 
spaced, and are separated by spaces of about the same diameter as themselves. 

None of the plates is distinctly subdivided, but each plate of the pairs is divided 
into three bands, the anterior and posterior darker, and the one between lighter, with 
smaller dots. The first median is far separated from the shoulder plate, and near the 
first pair. The third median is somewhat obscure, being little more than a dotted 
area with obscure margins, close to the second pair. 

The claws are unusually large, being about one-tenth of the length of the body, 
but there are species in which they are still longer (relatively). There is a palp near 
the base of the fourth leg, and a very short spine on the first leg. 



TARDIGKADA: PACIFIC ISLANDS 158 

E. viridis has been known for many years. The first specimens seen were empty 
skins, and it was considered unsafe to put any importance on the colour of the dead 
animals. The colour alone could not be considered of specific value, as it might be 
due to disease, and could only be regarded as having any weight if the animal 
showed other peculiarities. These we find in the nature of the dots, the very short 
head setae, and the very large claws. 

The animal was first found alive in Hawaii, and it was seen that the plates had 
the distinctive colour in life, and possessed the other peculiarities which had been 
noticed in the green skins previously observed. 

It appears, then, to be a good species, and the olive colour seems to be constant 
and characteristic. Disregarding the green colour there is no species with which it 
could be united. There are but few species having V. and VI. joined, and no 
processes after seta a. E. wendti and E. reticulatus have the seta a very long, and 
the latter has the surface reticulate and the spine on the front leg very long. 
E, arctomys has no fringe. E. kerguelensis and E. sylvamis have very fine dots on 
the plates. E. macronyx (38), which has similar large claws, has the surface finely 
punctate, and there are no barbs on the claws. The other species of the group differ 
conspicuously, and need not be compared. 

Habitat. Among moss from the bush near the City of Honolulu, Island of 
Oahu, Sandwich Islands. Previously found at the margins of two Scottish lochs 
(Ness and Morar). 

The dots on the plates vary somewhat in size, though always large ; the figure 
shows them of the largest size. 

Echiniscus spinulosus, Doyere (2) (Plate XIX. Fig. 38) 

Although differing from Doyere's species in one important point, viz., that the two 
dorsal processes, over c and d, are small spicules instead of long spines, I do not 
feel justified in making this Hawaiian form a distinct species. The dorsal processes 
vary more than any others in relative size. In a large series of specimens of 
E. duboisi, a species closely related to this one, and like it having only spines on the 
body, I have seen the dorsal processes vary from spicules to long spines. As the 
Hawaiian examples have some distinct characters, not noticed in Doyere's description, 
a full description is here given. These are in the surface texture of the plates, and 
have been insufficiently attended to in descriptions of species. 

Description. Size moderate, length 250ft exclusive of the legs. Plates nine, two 
pairs, two median. The surface texture is very striking. The dots appear very 
distinctly as perforations, some large and some small, with irregular wide spaces 
between. On each plate of the pairs the dots are confined to two areas, separated 
by a plain band on which there are no dots. The anterior area, along the border of 
the plate, is narrow, the posterior area occupies more than half the plate. 

BK1T. ANTARCT. EXPED. 1907-9. VOL. I. X 



154 J. MURRAY 

In front of the second median plate, from which it is separated by a plain band, 
and close to the edge of the first pair, there is a narrow dotted band. There is a 
similar band behind the second pair of plates, which I have not considered as a 
median plate. These bands, and the separate dotted areas on the paired plates, 
recall the intercalary plates of Bichters' E. scrqfa and E. quadrispinosa (26) and 
are doubtless of similar value. 

The lumbar plate is trefoliate and faceted, having four principal panels of which 
the posterior one is obscurely divided in two. The spines of the fringe on the fourth 
leg are short triangles. The decurved spine of the inner claws is somewhat large 
and far from the base of the claw. 

The oval eggs are about 80 M long, by 60 f* wide. Two or three are laid in 
the skin. 

Australian varieties of E. duboisi have precisely the same surface texture, and 
the same subdivision of the dotted area of the paired plates. This confirms the 
close affinity of the two species. Such peculiarities of surface texture must not be 
given too much weight in separating new forms from old, as there can be little doubt 
that they have been generally overlooked. 

Echiniscus perarmatus, Murray ? (20) 

Identified from a mutilated skin, having none of the spines or setse. The plates 
with dots of two kinds, the broad plain band at the posterior margin of the paired 
plates, and the papillae showing on the very edge of the lumbar and median plates, 
are all characters unknown in any other species, or at any rate not combined in 
any species. 

Echiniscus, sp. ? (Plate XIX. Fig. 37) 

A small animal, probably immature, which could not be identified. Several 
examples were seen. 

Description. Length 125 p, exclusive of the legs. Plates nine, two pairs, two 
median ; dots fine, regular, nature doubtful. Lateral processes a and e, a a seta 
of 50 , e a curved spine of 12 n, sometimes lacking. Dorsal process over d a short 
curved spine of 6 /*. On fourth leg a fringe of small blunt teeth. All claws without 
barbs. Lumbar plate trefoliate. 

It is needless to make any comparison with other species, as the animal appears to 
be young and incompletely developed. 

Genus Milnesium, Doyere (2) 
Milnesium tardigradum, Doyere (2) 
The branched claws had, some three, some two points. 



TARDIGRADA: PACIFIC ISLANDS 155 

Genus Macrobiotus, Schultze (42) 

A : SPECIES HAVING KOUGH EGGS 

Macrobiotus intermedium, Plate (23) 

Identified from an egg which contained a well-developed young. The egg had 
the typical top-shaped processes. It was, however, not spherical but shortly oval, 
measuring 60 by 50 n over the spines. The pharnyx contained three quadrate 
nuts in each row, the first apparently united to the gullet, but distinct from the 
apophysis. 

Another elliptical spiny egg occurred in Hawaii (Plate XIX. Fig. 416). 

Macrobiotus crassidens, Murray (20) 

As the egg was not found there is some little doubt about this record. It is 
distinguished from the preceding by the greater relative breadth of the nuts in the 
pharynx, which are even broader than long. 

Macrobiotus oberhduseri, Doyere (2) 

The Hawaiian form of this species was papillose all over, as in Central African 
examples, but the papillae were smaller. The bands of colour were faint. The 
pharynx contained two nuts in each row. 

It is likely that this all-papillose species will prove to be distinct from Doyere's 
type. The eggs of the various forms of this species are not yet sufficiently known. 

Macrobiotus occidentals (?), Murray (Plate XIX. Figs. 39a-39c) 

This Macrobiotus, differing in some respects from the type of M. occidentalis, and 
probably a distinct insular race or species, cannot be assigned its final place till the 
egg is known. 

Description. Small, hyaline or very pale yellow. Length 300 /*. Skin thin, 
dotted. In all the examples seen there was never the double skin, enclosing a clear 
fluid filled with thin hyaline rectangular plates, as in the type. Teeth small, not 
widely spreading, abruptly bent outwards beyond the middle. Gullet narrow ; 
pharynx with apophysis and two short rods in each row, the first (next the gullet) 
more than twice as long as the second, which is shortly oval ; a small comma. Small 
black eyes. Fat-cells very large, 10 to 15 ^ in diameter, hyaline. 

A large (old) example measured 450 /j. in length, and was pigmented like 
M. hufelandii. The claws are slender, and united less than half-way. 

The points of difference from the type are : the smaller size, lack of colour in the 
fat-cells, lack of double skin, abruptlybent teeth, and comma in the pharynx. These 
are points of unequal value, in the aggregate of considerable importance. The pale 



156 J. MURRAY 

colour might be associated with youth, but even the eggs of the type have dull 
reddish contents. The double skin, with enclosed fluid and plates, is not regarded as 
a specific character, although all the Canadian examples had it. It is possibly 
pathological. The comma is also of uncertain value, though I believe generally 
constant in a species. It remains for the discovery of the egg to decide the position of 
the form. Although pretty abundant no eggs were found which might belong to it. 
An almost identical form was found in Australia. 

Macrobiotus, sp. ? (Plate XIX. Figs. 41a, 416) 

An elliptical egg, with processes of the hufelandi type, and containing a young 
with well-grown pharynx. The egg measures 90 M by 65 M over the processes, which 
are about 6 /u in length. The processes are of the typical hufelandi form, but many 
of them are divided at the ends, as in furcigir, orcadensis, and a variety of 
intermedius. 

Teeth slender, curved outwards about the middle ; gullet of moderate width ; 
pharynx shortly oval, with apophysis and two rods in each row, the first twice as long 
as the second. The claws were not seen. 

B : EGGS UNKNOWN. 
MacroUotus, sp. ? (Plate XIX. Figs. 42, 426). 

A hyaline form, of which the egg is unknown. The claws are of the hufelandi 
type, but they are only joined for a short way above the base, and one of each pair 
is much shorter than the other. 

The gullet is narrower than in hufelandii and has a prominent rim at its end in the 
pharynx. The pharynx is shortly oval and has two rods and a comma in each row of 
thickenings. The first rod is more than twice as long as the second, and has a 
rounded projection near its base. The comma is very small. The apophysis was not 
observed. Eyes present. Length 350 M- 

Without the egg the species cannot be identified. It has some affinity with 
hufelandii, but has the claws united for a shorter distance, narrower gullet, and weaker 
teeth. The form of the first rod seems distinctive. 

Macrobiotus, sp. ? (Plate XIX. Figs 43, 436). 

Pharynx like that of M. virgatus (p. 173), but with a comma. Claws different, 
united at base only, unequal. 

REMARKS ON THE TARDIGRADE FAUNA or THE PACIFIC ISLANDS 

The interest of the Tardigrade fauna of these islands is chiefly geographical. 
They have not yet yielded any peculiar insular -forms in this group, although in some 
cases varieties may indicate peculiar insular races. 



TARDIGRADA: PACIFIC ISLANDS 157 

Our knowledge is as yet far too scanty to permit of any useful discussion even of 
geographical distribution. All the collections have been hurriedly made in the most 
unlikely localities, viz., in the immediate neighbourhood of the various ports of call. 
The entire interior of the islands, with all their variety of climate and physical features, 
is untouched. 

No doubt the right kind of work would result in greatly extending the number of 
species. The great apparent difference in the productiveness of Fiji and Hawaii is 
doubtless due to nothing else than the fact that there was no suitable collecting- 
ground within reach at Fiji. 

All that can at present be done is to indicate the general distribution of the few 
species yet collected. 

In the following table I have included Professor Richters' records for Samoa and 
the Hawaiian group (37). These include one species (M. samoanus) which we did not 
get, and the only species which is not known anywhere else than in the Pacific 
Islands. It exhibits the distribution in four columns for the islands, and nine for the 
rest of the world. 



DISTRIBUTION : LOCAL AND GENERAL 



Pacific Islands 















Q 


















b 



i-l 

H 

IH 


SAMOA. 


OAHU. 


HAWAII. 


USTRALIA. 


w ZEALAN 


EUROPE. 


en 


AFRICA. 


. AMERICA 


AMERICA. 


ARCTJC. 


NTARCTIC. 




^ 








^3 


m 








^ 


oj 




<3< 














X 
















Echiniscus mutabilis 








X 





X 


X 


X 


X 


X 





. 


X 





E. noi-cezeelandia 








X 





X 


X 























E. arctomys (?) * 








X 





X 





X 


X 


X 








X 


X 


E. intermedius 








X 





X 














X 





- 





E. viridis 








X 











X 




















E. spinulosus . 








X 











X 














X 





E. perarmatus 








X 

















X 














Milnesium tardiyrudmn 








X 





X 


X 


X 


X 


X 


X 


X 


X 


X 


Jfacrobiottis hufelandii 








X 


X 


X 


X 


X 


X 


X 


X 


X 


X 





M. areolatus . 


X 











X 





X 


X 


X 


X 





X 


- 


M, occidentalis 








X 





X 














X 











M. intermedius 








X 





X 


X 


X 


X 


X 


X 


X 


X 





M. crassidens . 








X 





X 











X 














M. oberfuiuseri 

J.YL > 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 














a 


a 








P 




H 




*i 








<i 


Jz; 


K 




o 




S 


CO 




J 





ri 




J 


M 

3 


^ 


J 


o" 






j 


E 


r 2 





03 


M 


*"'" 





9 




OJ 




o 




O 
O 


* 




O 





B 

a 


-O 


*3* 


3 




H 


g 

tsi 


D 


o 




1 







o 




I 




tJ 


^N 











^ 


H 




<1 


M 


-^ ^ 


o 




H 




^ 


s> 


fe 


5 


<IJ 










3 




02 








^ 


^ 


o 




* 
























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 

eB 

<! 


AFRICA. 


ARCTIC. 


ISLANDS. 


Echiniscus mutabilis, Murray .... 





X 


X 


X 








X 


X 


X 


X 





E. novcezedandice, Richters .... 





X 


X 


X 























E. pulcher, sp. n. . 








X 


























- E. arctomys, Ehrenberg ? * 


X 





X 


X 








X 


X 


X 


X 


- 


E. kerguelensis, Richters ..... 








X 























X 


K. sylvanus, sp. n. 














X 




















E. viridis, sp. n. ...... 











X 








X 














E. tessellatus, sp. n. . 








X 


I 























E. gladiator, Murray ..... 





X 








X 





X 






















x 


x 


x 




















E. canadensis, sp. n. 












X 




















E. bisetosus, Heinis ...... 














X 





X 














E. spinulosus, Doyere ..... 











x 








x 







x 





E. spiniger, Richters ..... 








X 










X 













K. duboisi, Richters ...... 








X 














X 


X 








K. perarmatvA, Murray ..... 





. 


. 


X 














X 








E. blumi, Richters ...... 








X 











X 








X 





E. velaminis, sp. n. . 





X 





























E. oihonnce, Richters ..... 








X 





X 





X 








X 





Oredla niollis, gen. n., sp. n. . . 








X 

























-Milnesium tardigradum, Doyere 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


X 


Macrobiotus hufelcmdii, Schultze 





X 


X 


X 


X 


X 


X 


X 


X 


X 


X 




x 






















M. echinogenitus, Richters .... 


X 


X 


X 





X 


_ 


X 


X 


X 


X 


X 


M. areolatus, sp. n. . 








X 


X 


X 





X 


X 


X 


X 


X 


M. harmsworthi, Murray ..... 





X 


X 




X 





X 






X 




M. montanus, sp. n. . 





X 











































































M. occidentalis^ sp. n. 


_ 




X 


X 


X 














M. intermedius, Plate ..... 





X 


X 


X 


X 


x 


X 


X 


x 


x 


x 


M. crassidens, Murray ..... 







X 


X 










x 






M. aculeatus, sp. n. . 






x 




















5 
























s 












X 


X 


X 


X 


X 


M. arcticu,s, Murray 


~*S 


X 


X 






X 


X 


X 




X 




M. tuberculatus, Plate ..... 


x / 




X 




X 

x 




x 




X 

x 


X 

x 




M. nodosus, Murray ...... 




x 




x 










x 






M. sattleri, Richters . 




























X 


X 




x 




X 


X 






X 


M. ornatus, Richters 




X 


X 








X 










M. rubens, Murray ..... 
M. annulatus, Murray .... 





X 

x 


X 











X 

x 


X 


X 


X 


X 


M. canadensis, sp. n. . , 
























M. augusti, Murray ..... 






x 








x 






x 




M. virgatus, sp. n. 
























Diphascon chilenense, Plate .... 
D. alpinum, Murray . 
D. scoticum, Murray . 
D. canadense, sp. n. . 


X 
X 
X 


X 
X 
X 


_ 

X 
X 


X 


X 
X 
X 
X 


X 
X 


X 
X 
X 


X 





X 
X 
X 


X 



























Twenty -three species not identified 1 2 of Echiniscus, 10 of Macrobiotus, and 1 doubtful, MacrMntHK or 

Dipfuwoon, Total, 73 species. 
* All these records of E. arctomys, except for Australia, are wrong (see p. 126 and footnote). 






TARDIGRADA 181 

described as new species, and 23 could not be identified. Of the 50 named species 
26 were got in one only of the countries visited ; 14 of these 26 are known in other 
parts of the world, while 12 are as yet unknown except in the one locality for each 
here recorded. These last are, of course, among the new species collected by the 
Expedition. Five of the new species were found in more than one country, or were 
previously known elsewhere, though they had not been described. 

Australia has most species, viz., 38 (31 identified), Canada comes next with 31 
(23 identified), New Zealand has 25 (21 identified), the Pacific Islands 18 (15 
identified), and the Antarctic 5 (4 identified). 

Canada has 7 of the new species, Australia 6 (and 1 new genus), New Zealand 2, 
the Pacific Islands 1, and the Antarctic 1. 

It is noticeable how much richer the continental areas are than the islands, both 
in the number of species and the proportion of peculiar species. The isolation of 
New Zealand, Fiji, and Hawaii does not appear to have led to the development 
of many new forms. 

As the Tardigrada of Europe are much more fully known than those of any other 
part of the world, the list of European species may be used as a standard, and it may 
be instructive to compare the lists of species from the various countries with it, and to 
note how much they have in common. Canada has 16 species in common with 
Europe, Australia 17 species, New Zealand 16 species, Pacific Islands 9 species, and 
Antarctic 3 species. Taking the ratios of these numbers to the totals for each 
country, it appears that subtropical Australia has in its Tardigrade Fauna nearly 
as much correspondence with that of Europe as has temperate Canada, where the 
climatic conditions are so similar to those of northern Europe. New Zealand and the 
Antarctic have more correspondence than either. For what those figures are worth, 
New Zealand is most like Europe, the Antarctic comes next, then Canada, Australia, 
and last of all the Pacific Islands. But there is little to choose between them, and 
the figures are not worth much, as they would be liable to change whenever further 
work is done in these countries. 

The peculiarity of the Australian Tardigrade Fauna is greater than we would 
suppose from a mere consideration of the number of peculiar species. Not only is there 
a distinct generic type (Oreella), but most of the species (E. pulcher, E. tessellatus, 
E. interniedius), are conspicuously different from their nearest relatives. Even 
M. aculeatus, though differing from the African M. crassidens mainly by one external 
character, is of great interest. The six spines, occupying definite positions as they 
do on the segments which bear the second, third, and fourth pairs of legs, may 
possibly be the homologues of the dorsal processes of Echiniscus. If this were 
the case the occurrence of this species, and of the genus Oreella, would be of 
great importance in elucidating the affinities of fhe genera Macrobiotus and 
Echiniscus, 

The existence of these various peculiar forms in Australia serves to indicate that 
the Australian native Tardigrada are partly at least of very ancient origin, although 



182 J. MURRAY 

on account of the comparative facility of immigration, such facts have less weight in 
this group than in the Vertebrata. 

The further study of the Tardigrada of continental and insular areas, in which 
cognisance would be taken of all known species, is alluring, but beyond the scope of 
this work. 

Natural History. There is in this paper, unfortunately, too little study of pure 
biology, or the natural history of the water-bears, except in the wide sense that the 
form of every organism is a manifestation of life. In this sense, every trifling 
variation of form could throw some light on the nature of the living force which 
produces it, although we may be unable to trace its meaning. 

The conditions under which the work was done prevented much attention being 
given to the study of living animals. Nearly all of the species were seen alive, so 
that the material for the investigation of life-histories was available, yet we know 
nothing as to such points as the duration of life, the time taken to grow up, the 
changes that occur during growth, the relations of the sexes, the time required for 
the eggs to hatch, and a host of others. 

The observation of the living animals so closely as to elucidate any of these 
points, while fascinating to the naturalist, is too tedious, or if not tedious yet takes 
far too long, to be compatible with the compilation of a series of reports on the 
collections of an expedition. 

There is no suggestion made as to the meaning of the peculiarities of species in 
their relation to their surroundings. The value of specific peculiarities to their 
possessors is generally very obscure, so much so that in an earlier chapter dealing 
with the Value of Species (p. 92) I have suggested that many of the specific 
characters are " fortuitous." Fortuitous in the ordinary sense they cannot be, but 
they may be so described in the restricted sense that they are of no definite use to 
the species. They may have been produced by the interaction of the conditions and 
the living force, without leading to more complete adaptation to the conditions. 
This must be understood as applying only to certain characters, since constant 
adaptation must have gone on as well. 

We can understand or guess at the reasons why it is advantageous to some 
species to deposit smooth eggs in the skin at the moult, while others lay rough eggs 
without the protection of the skin we can imagine benefits from the strange 
simplification and encystment of water-bears. These things are obviously important, 
whether we fully understand them or not. 

There are, however, many little peculiarities of outward form, those things which 
supply the specific characters generally, in which it is difficult to see any advantage. 
They may, of course, have a value which is hidden from our eyes. Such are the 
spines and setee of Echiniscus. It seems a reasonable supposition that such spines 
are protective : they may deter an enemy from making a meal of their possessor. 
But there are species with the processes quite soft and weak, such as the acicular 



TARDIGRADA 183 

spines of E. africanus, and these cannot be protective ; while the commonest of all 
species have no spines. 

In one direction the study of the variations of form among the Tardigrada is 
important, although we may not see the causes of the variation or its value to the 
animals. Through the various forms we can learn the affinities of the species, and 
we may thus gain some knowledge of the history of the development of the group. 
Thus the Tardigrada are brought into geological history, although their remains may 
be scarcely known in the strata of the earth. 

It is an interesting fact in itself that so many Tardigrada have been obtained 
alive from dried mosses. Although a few were studied while the mosses were fresh, 
the great majority of the fifty species were got from mosses which had been dried for 
a longer or shorter period, varying from a month or two to nearly three years. 
This brings out strongly the power possessed by habitual moss-dwellers of sur- 
viving long periods of drought. After being kept for a year perfectly dry the 
number of animals which revive on being moistened seems no less than when the 
moss has only been a short time dry. After a year the vitality rapidly diminishes 
after three years a very small proportion of the animals survive. The duration 
of their vitality is greatly affected by conditions. If there is a suspicion of 
moisture, if enough to produce mildew, or even merely to cause a musty smell, 
they will not live long. 

In the Antarctic some experiments were made in order to test the limits to the 
vitality of water-bears. These are detailed briefly under the account of Macrobiotus 
arcticus. 

BIBLIOGRAPHICAL LIST OF WORKS REFERRED TO IN THE TEXT 

Several others, which came too late to be inserted in their proper places, will be found in footnotes. 
In the text the references to this list are made in figures in heavy type, enclosed in parentheses. 

1. CARLZON, 0., " Schwedische Tardigraden," Zool. Anzeig., Bd. 34, 1909, p. 137. 

2. DoYiRE, M. T., "Memoire sur les Tardigrades," Ann. des sci. nat., Ser. 2, Tom. 14, 1840, p. 269. 

3. EHBENBERG, C. G., " Diagnoses novarum formarum," Verh. K. Akad. Wiss. Berl., 1853, p. 530. 

4. " Mikrogeologie," Atlas, Taf. 35u, 1854. 

5. " Mikroscopischen Lebens in bis 20,000 Fuss Alpenhb'he," Abhand. K. Akad. Wiss 

Berl. (aus dem Jahre 1858), 1859, p. 429. 

6. HAY, W. P., "A Bear-Animal renamed," Proc. Biol. Soc. Washington, xix., 1907, p. 46. 

7. HEINIS, Fr., "Tardigraden der Schweiz," Zool. Anzeig., Bd. 32, 1908, p. 633. 

8. "Moosfauna der kanarischen Inseln," Zool. Anzeig., Bd. 33, 1908, p. 711. 

9. ,. ,. " Moosbewohnenden Rhizopoden, Rotatorien, und Tardigraden," Arch, fur Ilydrobiol. 

und Planktonkunde, Bd. 5, 1910. 

10. HUTTON, F. W., "Index Faunae Novae Zealandia:," London, 1904. 

11. MUKEAY, J., " Tardigrada of the Forth Valley," Ann. Scot. Nat. Hist. 1905, p. 160. 

12. "The Tardigrada of the Scottish Lochs," Trans. Roy. Soc. Edin., vol. xli., 1905, p. 677. 

13. "Tardigrada of the Forth Valley," 2nd paper, Ann. Scot. Nat. Hist., 1906, p. 214. 

14. "Scottish Alpine Tardigrada," Ann. Scot. Nat. Hist., 1906, p. 25. 

15- ,, " Tardigrada of the South Orkneys," Trans. Roy. Soc. Edin., vol. xlv., 1906, p. 323. 



184 J. MURRAY 

16. MURRAY, J. "The Encystment of Macrobiotus," The Zoologist, 1907, p. 1. 

17. " Some Tardigrada of the Sikkim Himalaya," Journ. Roy. Micr. Sac., 1907, p. 20!). 

18. ,, "Scottish Tardigrada, collected by the Lake Survey," Trans. Roy. Soc. Edin., vol. xlv., 

1907, p. 041. 

19. "Arctic Tardigrada, collected by Wm. S. Bruce," Trans. Roy. Soc. Edin., vol. xlv., 

1907, p. GC9. 

20. "Some South African Tardigrada," Journ. Roy. Micr. Soc., 1907, p. 515. 

21. PACKARD, A. S., " Discovery of a Tardigrade," Amer. Naturalist, vol. vii., 1873, p. 740. 

22. PERTY, M., "Die Familie Xenomorphidse," Okens Ms (Jahrg. 1834), p. 1241, 1835. 

23. PLATE, L. H., " Naturgeschichte der Tardigraden," Zool. Jahrb., Abt. f. Anat. Bd. 3, 1888, p. 47. 

24. RICHTERS, F., "Fauna der Umgegend von Frankfurt-a.-M.," Her. Senckbg. Naturf. Ges., 1900. 

25. ,, ,, "Neue Moosbewohner," Ber. Senckbg. Naturf. Ges., 1902. 

26. " Fauna der Umgebung von Frankfurt-a.-M.," Ber. Senckbg. Naturf. Ges., 1902. 

27. " Nordische Tardigraden," Zool. Anzeig., Bd. 27, 1903, p. 168. 

28. " Verbreitung der Tardigraden," Zool. Anzeig., Bd. 28, 1904, p. 347. 

29. ,, "Echiniscus conifer," Ber. Senckbg. Naturf. Ges., 1904, p. 73. 

30. " Islandische Tardigraden," Zool. Anseig., Bd. 28, 1904, p. 373. 

31. " Antarktische Moosfauna," Verh. deutsch. Zool. Ges., 1904, p. 236. 

32. "DieEier der Tardigraden," Ber. Senckbg. Naturf. Ges., 1904, p. 59. 

33. " Arktische Tardigraden," Fauna Arctica, Bd. 3, Jena 1904, p. 495. 

34. ,, "Zwei neue Echiniscus- Arten," Zool. Anzeig., Bd. 31, 1907, p. 197. 

35. ,, ,, "Fauna der Moosrasen des Gaussbergs, &c.," Deut. Siidpol. Exped., 1901-3, Bd. 9 

Zool. 1907, p. 261. 

36. " Antarktische Tardigraden," Zool. Anzeiy., Bd. 31, 1907, p. 915. 

37. ,, " Moosfauna Australians, &c.," Zool. Jahrb., Bd. 26, Abt. fiir Syst. 1908, p. 196. 

38. "Moosbewohner," Wiss. Ergebn. Schwed. siidjwl. Exped., 1901-3, 1908. 

39. " Moosfauna-Studien," Ber. Senckbg. Naturf. Ges., 1908, p. 14. 

40. "Tardigraden unter 77 S. Er.," Zool. Anzeig., Bd." 34, 1909, p. 604. 

41. SCHAUDINN, F., " Die Tardigraden," Fauna Arctica, Bd. 2, 1901, p. 187. 

42. SCHULTZE, C. A. S., "Macrobiotus hufelandii," Okens Isis., 1834, p. 708. 

43. " Echiniscus bellermanni," Berlin, 1840. 

44. "Echiniscus creplini," Gryphire, 1861. 

45. SCOUKFIELD, D. J., "Non-Marine Fauna of Spitsbergen." Part I. Rhizopoda, Tardigrada 

Entomostraca, &c., Proc. Zool. Soc. London, 1897, p. 790. 

46. WHITELEGGE, T., " Invertebrate Fauna of Port Jackson and neighbourhood," Journ, and Proc. Roy. 

Soc.N.S.W., 1889, p. 163. 



TAUDKiKADA 



1.S.3 



INDEX TO SPECIES 



JJiphascon alpiniim, Murray, 100, 119, 

175 

D. canadense, sp. n., 176 
D. chilenense, Plate, 119, 143, 175 

D. scoticum, Murray, 119, 143, 176 
Echiniscus arctomys, Ehrenberg, 128 

E. bisetosus, Heinis, 164 
E. blumi, Richters, 132 
E. canadensis, sp. n., 162 
E. duboisi, Richters, 131 

E. gladiator, Murray, 111, 160 

E. intermedius, sp. n., 129, 151, 161 

E. kerguelensis, Richters, 128 

E. mutabilis, Murray, 110, 126, 151 

E. novc'zeelandia;, Richters, 110, 126, 151 

A', oihonnii', Riolitcrs, 132, 103 

E. perarmatus, Murray, 154 

K. pidcfter, sp. n., 127 

E, spiniyer, Richtei's, 131 

E. spinulosus, Doyere, 153 

E. sylvanus, sp. n., 160 

E. tessellatus, sp. n., 129 

E. vela-minis, sp. n., 112 

E. viridis, sp. n., 152 

Macrobiotus aculeatus, &p n., 139 

J/. annulatus, Murray, 118 



M. arctious, Murray, 96, 118, 140, 171 
M. areolatus, sp. n., 137, 148, 167 
M. augusti Murray, 141 
M. canadensis, sp. n., 171 
M. crassidens, Murray, 139, 155 
~~M~dispar, Murray, 117, 140 
M. eckinoyenitus, Richters, 115, 137, 167 
M.furtiger, Murray, 114 
M. harmsworthi, Murray, 115, 137, 169 
M. hufdandii, Schultze, 114, 137, 166 
M. hufelandioides, sp. n., 138 
M. intermedius, Plate, 117, 139, 155, 171 
M. montanus, sp. n., 116 
M. iwdosiis, Murray, 118, 148 
M. oberhfiuseri, Doyere, 99, 155, 171 
M. occidentalis, sp. n., 139, 155, 169 
M. ornatus, Richters, 118 
M.papittifer, Murray, 118, 141 
M. polaris, sp. n., 98 
M. rubens, Murray, 141 
M. sattleri, Richters, 118, 141, 173 
M. tuberculatus, Plate, 173 
M. virgatus, sp. n., 142, 173 

Milnesium tardiijradum, Doyere, 114, 137, 

154, 166 
Oreella mollis, gen. n. ; sp. n , 135 



B1UT. AXTAUCT. EXPED. 1907-D. VOL. I. 



1 15 



PLATE XI V 



PLATE XIV 

ANTARCTIC TARDIGRADA 

PAGE 

FIGURE la. Macrolnotm polaris, sp. n., adult . . . . x 300 98 

FIGURE 16. The same, teeth and pharynx 

FIGURE lo. The same, young emerging from the egg x 300 

FIGURE Id. The same, variety of the egg ..... x 300 

FIGURE le. The same, claws 

FIGURE 2. Macrobiotus arcticus, Murray, adult . . . . x 300 96 

FIGURE 26. The same, teeth and pharynx of adult 

FIGURE 2e. The same, thick-shelled egg x 300 

FIGURE 2d. The same, claws of adult 

FIGURE 2e. The same, young just hatched ..... x 300 

FIGURE 2f. The same, teeth and pharynx of young 

FIGURE 3. Diphascon alpinum, Murray, Antarctic variety . . x 300 100 

FIGURE 4. Diphascon or Macrobiotus (?). A doubtful intermediate 100 

form. Teeth and pharynx, showing shortly elongated 
gullet. The claws are like Fig. 2d on this plate. 



Brit Antarct Exped 1907-9. Vol. I. Plate XIV 

MURRAY ANTARCTIC TARDIGRADA. 












i POLARIS, sp.n. 2.M.ARCTICUSMURRAY. 3. D IPHASC N ALPINUM MURRAY 4.DlPHASCONor MACROBIOTUS. 

Murray del. ad nat 



PLATE XV 



PLATE XV 

TARDIGRADA OF NEW ZEALAND 

PAGE 

FIGURE 5. Ecliiniscus novcezeelandice, Richters x 300J 

I 126 

FIGURE 6. Echiniscus velaminis, sp. n. ..... x 300 112 

FIGURE 7. Echiniscus, sp. ? . . . . . . . x 300 113 

FIGURE 8. Echiniscus, sp. ? . x 300 113 

FIGURE 9. Echiniscus, sp. ? x300113 

FIGURE 10a. Macrobiotus montanns, sp. n x 300 11 G 

FIGURE 10&. The same, claws 

FIGURE lOc. The same, teeth and pharynx 

FIGURE lOd. The same, egg x 300 

FIGURE lla. Diphascon chilenense, Plate ? teeth and pharynx . 119 

FIGURE 116. The same, claws 

FIGURE 12. Macrobiotus, sp. ? Macquarie Islands ... 118 



Brit Antaret Exped 1907-9. Vol I. Plate XV 

TARDIGRADA OF NEW ZEALAND 




5. ECHINISCUS NOVAEZEELANDIAE RlCHTERS. 6.-E VELAMINIS sp.n 7 8 &9. ECHINISCUS SPECIES. 7 

10. MACROBIOTUS MONTAKUS. spn. 1.2. MACROBIOTUS SPECIES' 11. DIPHAS CON CHILENENSE PLATE 

Murray del ad nat 



PLATE XVI 



PLATE XVI 

AUSTRALIAN TARDIGRADA 



PAGE 



FIGURE 13. Echiniscus kerguelensis, Richters x 300 128 

FIGURE 14. Echiniscus blumi, Richters x 300 132 

FIGURE 15. Echiniscus tessellatus, sp. n. ..... x 300 129 

FIGURE 16. Echiniscus, sp. ? x 300 135 

FIGURE 17. Echiniscus intermedius, sp. n., type with reticulate 

surface ; x 300 129 

FIGURE 18. Echiniscus, sp. ? x 300 133 

FIG. 14. E. blumi is marked to indicate the segments I. to VI. (Richters), and the 
lateral processes a, b, c, and d. Those four processes are seise, but e is lacking in this 
species. At the base of a is the " auricle." Segments V. and VI. having completely 
coalesced, the single segment resulting is referred to as V. + VI. The figure shows 
also a little sharp spine on the first leg, a blunt palp at the base of the fourth leg, 
the "fringe" of sharp teeth, and the " barbs " of the outer and inner claws. 



Brit Antarct Exped 1907-9. Vol I. Plate XVI 

MURRAY- AUSTRALIAN TARDIGRADA. 





17 





18 



13.ECHINISCUS KERGUELENSIS RlCHTERS. 14-. E.BLUMI RlCHTERS. 15. E. TESSELLATUS. sp n 

17. E INTERMEDIUS spn 16. & 18.ECHINTSCUS SPECIES.? 
J Murray del ad. naL 



PLATE XVII 



BRIT. ANTARCT. EXPED. 1907-9. VOL. I. 2 C 



PLATE XVII 

AUSTRALIAN TARDIGRADA 



PAGE 



FIGURE 19. Echiniscus duboisi, Richters, variety x 300 131 

FIGURE 20. The same, another variety x 300 132 

FIGURE 21. Echiniscus oihonnce, Richters ..... x 300 132 

FIGURE 22. Echiniscus, sp. ? x 300 134 

FIGURE 23. Echiniscus, sp. ? larva x 400 135 

FIGURE 24. Echiniscus spiniger, Richters, variety . . . x 300 131 

FIGURE 25. Echiniscus, sp. ? .... x 300 134 



Brit Antarct Exped 1907-9. 



Vol. I. Plate XVII 



AUSTRALIAN TARDTGRADA.. 




22. 



19.& 20. ECHINISCUSDUBOISI RlCHTERS. 21. E.OIHONNAE RlCHTERS. 24.E. SPINIGERRlCHTERS. 

22.23.k25. ECHINISCUS. SPECIES? 
J Murray del. ad nat 



PLATE XVIII 



PLATE XVIII 

AUSTRALIAN TARDIGRADA 

PAGE 

FIGUEE 26. Oreella, gen. n., mollis, sp. n x 300 135 

FIGURE 27a. Macrobiotus aculeatus, sp. n x 300 139 

FIGURE 27b. The same, teeth and pharynx 

FIGURE 27c. The same, egg . ... x 300 

FIGURE 27d. The same, claws 

FIGURE 27e. The same, variety with only two spines x 300 

FIGURE 28. Macrobiotus virgatus (?) 142 

FIGURE 29. Macrobiotus hufelandioides, sp. n. . . . 138 

FIGURE 29b. The same, three processes of the egg 

FIGURE 29c. The same, claws 

FIGURE 30. Macrobiotus echinogenitus (?) egg x 300 137 

FIGURE 31. Macrobiotus harmsworthi (?) egg x 300 137 

FIGURE 32. Macrobiotus, sp. ? foot, showing knob ... 142 

FIGURE 32&. The same, claws 

FIGURE 32c. The same, teeth and pharynx 

FIGURE 33a. Macrobiotus, sp. ? x 300 142 

FIGURE 336. The same, claws 

FIGURE 33c. The same, teeth and pharynx 

FIGURE 34. Echiniscus pidcher, sp. n x300127 

This figure is marked to indicate the segments I. to VI. (Richters), and the 
lateral processes a, b, c, d, and e. In this species a and e are setse and ?>, c, d are 
little cones. At the base of a is the " auricle" Segments V. and VI. are separate. 



Brit Antarct Exped 1907-9. 



AUSTRALIAN TARDIGRADA.. 



Vol. I. Plate XVHI 




33c. 



27e. 



26. ORBELLA . ge n MOLLIS n. sp . 27MACROBIOTUS ACULEATUS sp ,-> 28.M.viRCATUs MURRAY: ? 

29.M HUPLLANDIOlDESspn SO.M.ECHINO&ENITUSRlCHTERS. 31.M.HARMSWDRTHI MURRAY: 34-.ECHINTSCUSPULCHER.spn 

J. Murray del. ad nat. 32 ^3.MACRDBiOTrjs. SPECIES ? 



PLATE XIX 



PLATE XIX 

TARDIGRADA OF PACIFIC ISLANDS 

PAGE 

FIGURE 35. Echiniscus novc&zeelandice, Richters x 300 151 

FIGURE 36a. Echiniscus viridis, sp. n. x 300 152 

FIGURE 366. The same, inner claw 

FIGURE 36c. Echiniscus intermedius, Murray . . . . x 300 151 
FIGURE 36d The same, portion of reticulated surface, more highly 
magnified 

FIGURE 37. Echiniscus, sp. ? x 600 154 

FIGURE 38. Echiniscus spinulosus, Doyere, variety . . . x 300 153 

FIGURE 39a. Macrobiotus occidentalis, Murray x 300 155 

FIGURE 396. The same, teeth and pharynx 

FIGURE 39c. The same, claws 

FIGURE 40. Macrobiotus areolatus (?) egg . . . . . x 300 148 

FIGURE 4 la. Macrobiotus, sp. ? teeth and pharynx ... 156 

FIGURE 416. The same, elliptical egg ... x 300 

FIGURE 42. Macrobiotus, sp. ? teeth and pharynx ... 156 

FIGURE 426. The same, claws 

FIGURE 43a. Macrobiotus, sp. ? teeth and pharynx ... 148 

FIGURE 436. The same, claws 



Brit Antarct Exped 1907-9. 



Vol I. Plate XTK 



TARDIGRADA OF PACIFIC ISLANDS. 




35 





41 a. 



416. 




39c 




4-26. 



.38 



3 S.ECHINISCUS NOVAEZEELANDIAE RlCHTERS. 36 A E VIRIljlS spn. 36?E.nNTTERMEDJUS MlTRRAY 38E.SPINULOSUS DOYERE. 
37ECHINISCUS.sp? 39.MACROBIOTUS OCCIDENTALIS MURRAY 40.T043.MACB.OBIOTUS sp .? 

J Murray del. ad nat 



PLATE XX 



PLATE XX 

CANADIAN TARDIGRADA 

PAGE 

FIGURE 44. Echiniscus, sp. ? x 260 165 

FIGURE 45. Echiniscus, sp. ? x 260 166 

FIGURE 46. Echiniscus, sp. ? x 260 165 

FIGURE 47. Echiniscus canadensis, sp. n. . . . . . x 260 162 

FIGURE 48. Echiniscus oihonnce, Bichters ?..... x 260 163 

FIGURE 49. Echiniscus sylvanus, sp. n. ..... x 260 160 

FIGURE 50. Echiniscus bisetosus, Heinis ?..... x 260 164 

FIGURE 51. Echiniscus gladiator, Murray, one pair of plates . 160 

FIGURE 52e*. Echiniscus intermedius, Murray, American variety . x 260 161 

FIGURE 526. The same, larva with two claws x 260 



Brit Antarct Exped 1907-9. Vol. I. Plate XX 

MURRAY CANADIAN TARDIGRADA. 




47 ECHINISCUS CANADENSIS.sp n. 48. E. OIHON1LE HlCHTERS. 7 49.E.SYLVA2STUS.sp. n. 50. E. BISETOSUS HEINIS.? 

51. E GLADIATOR MURRAY 52. E.INTERMEDIUS MURRAY 44-. TO 6. ECHINISCUS SPECIES. ? 

J Murray del. ad nat 



PLATE XXI 



BRIT. ANTARCT. EXPED. 1907-9. VOL. I. 2 D 



PLATE XXI 

CANADIAN TARDIGRADA 

PAGE 

FIGURE 53. Macrobiotus areolatus, Murray . . x 260 167 

FIGURE 536. The same, teeth and pharynx 

FIGURE 53c The same, claws 

FIGURE 53d. The same, egg ..... x 260 

FIGURE 53e. The same, variety of egg ? ..... x 260 

FIGURE 54a. Macrobiotus occidentalis, sp. n. . . . x 260 169 

FIGURE 546. The same, teeth and pharynx 

FIGURE 54c. The same, egg x 260 

FIGURE 54c. The same, claws 

FIGURE 54e. The same, double skin with dots and plates 

FIGURE 55. Macrobiotus virgatus, sp. n. . . . . x 260 173 

FIGURE 55b. The same, teeth and pharynx 

FIGURE 55c. The same, claws 

FIGURE 56. Egg of unknown Macrobiotus .... x 260 175 

FIGURE 57. Egg of unknown Macrobiotus . . . . . x 260 175 

FIGURE 58. Egg of M. echinogenitus, variety x 260 167 

FIGURE 59. Egg like that of M. dispar, &c. x 260 175 

FIGURE 60. Macrobiotus, sp. ? teeth and pharynx ... 174 

FIGURE 61. Macrobiotus canadensis, sp. n. .... x 260 171 

FIGURE 616. The same, teeth and pharynx 

FIGURE 61c. The same, claws 

FIGURE 61d The same, egg x 260 

FIGURE 62. Diphascon canadense, sp. n x 260 ' 176 

FIGURE 626. The same, teeth, gullet, and pharnyx 
FIGURE 62r. The same, claws 



Brit Antarct Exped 1907-9. Vol I. Plate XXI 

MURRAY CANADIAN TAJ*DIGRADA. 




53.MACRQBIOTUS AB.EOLATTJS MURRAY 5 4-.M.OcCIDENTALIS.sp.n. 55.M.V1RGATUS sp n 58.M.ECHINOGENITUS RlCHTERS. 
61 M CANADENSIS.spn. 620lPHASCON CANADENSIS. 56. 57 59 &60 MACROBIOTUS SPECIES.? 

J Murray del. ad nat 



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 VI 

RHIZOPODES D'EAU DOUCE 

BY EUGENE PENARD, DR. ES SCIENCES 



(WITH TWO PLATES) 



LONDON 
PUBLISHED FOR THE EXPEDITION BY WILLIAM HEINEMANN 

21 BEDFORD STREET, W.C. 

1911 

ISSUED SEPARATELY, JANUARY 1!)11 
PRICE THREE SHILLINGS NET 



REPORTS ALREADY ISSUED 

VOL. I. PART I., ON COLLECTING AT CAFE ROYDS 

BY JAMES MURRAY PRICE Is. 6cl. 

VOL. I. PART II., ON MICROSCOPIC LIFE AT CAPE ROYDS 

BY JAMES MURRAY PRICE .is. Ocl. 

VOL. I. PART III., ANTARCTIC ROTIFERA 

BY JAMES MURRAY PRICE 5s. Od. 

VOL. I. PART IV., MUSCI 

BY JULES CARDOT PRICE Is. Od. 

VOL. I. PART V., TARDIGRADA 

BY JAMES MURRAY PRICE 10s. Od. 



PART VI 

SARCODINA 

RIIIZOPODES D'EAU DOUCE 
PAK EUGENE PENARD, DK. ES SCIENCES 

(AVEC LES PLANCHES XXII., XXIII.) 

LES collections que M^r. J. Murray a bien voulu m'envoyer, pour 1'examen des 
Rhizopodes d'eau douce rapportds par FExpedition antarctique du Nimrod, me 
sont parvenues sous la forme soit de paquets de mousses ou autres vegetaux a 
1'etat sec, soit de sediments conserves dans la formaline. 

En rendant compte aujourd'hui du resultat de mes recherches, il me semble 
utile d'envisager ces collections comme formant deux groupes speciaux, et necessitant 
par la une distinction en deux chapitres nettement separes. 

Le premier groupe, represente dans les collections par six flacons de re"sidu 
a la formaline et cinq petits paquets de mousses rases, minuscules, feutrees, frag- 
mentaires et reduites pour ainsi dire a leur plus simple expression, concerne les 
terres antarctiques proprement dites, File Eoss avec le Cap Royds et ses petits lacs, 
Hut Point ou dans une expedition precedente hiverna la Discovery, et les 
Moraines echouees du continent. 

Avec le second groupe, nous aurons a nous occuper des Rhizopodes recueillis 
lors du voyage de retour, et qui me sont arrives sous la forme de dix-neuf paquets 
de mousses seches et de quinze flacons renferrnant du sddiment. 

Mais ce second chapitre, a son tour, comporte un certain nombre de sub- 
divisions : Ties meridionales du Pacifique, Nouvelle Zelande, Australie, lies du 
Pacifique en general, Vancouver et le Canada. 

Un tableau general des especes rencontrees aura son utilite, en permettant un 
coup d'ccil d'ensemble sur la distribution des especes. Ces dernieres, enfin, seront 
reprises une k une et donneront lieu a quelques observations d'interet general. 



BIUT. ANTARCT. EXPED, 1907-9. VOL. I. PAKT 6, ISSUED SEPAEATELY 1911 

/3 



204 E. PENARD 



CHAPITRE I 

ANTARCTIQUE 
1. Cape Royds, mare a sec (dried tarn) 

Le materiel recolte dans cette localite, en Avril 1908, par Mr. J. Murray, 
consistait exclusivement en fragments jaunatres de " weed," de cette matiere 
organique que les explorateurs ont trouvt$e en abondance dans les environs du Cap 
Royds, et sur la nature de laquelle on n'est pas encore au clair. 

Aucun rhizopode ne s'est montre dans cette " herbe," sauf peut-etre une Corycia, 
genre auquel paraissaient devoir se rapporter deux enveloppes vides et en mauvais etat. 

2. Cape Royds, mousses recoltees par J. Murray, le 26 Janv. 1909 

Ces quelques petits fragments d'une mousse courte et feutree se sont trouvds 
renfermer, avec une abondance relative, la Corycia flava de GREEFF, representee ici 
par une forme un peu speciale, de faible taille, et dont 1'enveloppe, d'une teinte claire 
et d'une transparence toute particuliere, a permis des observations biologiques qui 
n'avaient pu 6tre faites auparavant. 1 Ces rhizopodes, en effet, etaient vivants, et se 
sont montres tels apres un temps relativement tres court, une lieure ou deux d'immer- 
sion ; il m'a ete possible de les conserver plus de huit jours. 

Outre cette. espece, les mousses du Cap Iloyds ont fourni : 

Euglyplia laevis, plusieurs exemplaires, coquilles vides. 

Assulina muscorum, une seule enveloppe vide. 

Centropyxis aculeata, une petite enveloppe vide, de 95 M de diametre, et qui 
se rapportait a cette forme decrite par CASH comme var, spinosa, particuliere aux 
sphagnum et aux mousses. 

Arcella arenaria, une enveloppe vide. 

Heleopera petricola, une coquille vide, tres vieille et en mauvais etat, mais qui 
montrait bien les caracteres de 1'espece, et revetait encore cette teinte Idgerement 
violette que 1'on observe frequemment dans cet organisme. 

Dans cette station du Cap Iloyds, se trouvait enfin certainement Trinema 
enchelys, & en juger d'apres un croquis que Mr. Murray m'a envoye, et qui ne 
peut laisser aucun doute sur 1'espece. 

1 Ces observations trouveront leur place dans un m^moire qui sera public prochainement, dans la Revue 
Suisse de Zoologie (Tome 19, 1911). 



RHIZOPODES D'EAU DOUCE 205 

3. Substance organique, remplissant des poches (Vegetation in 
pockets, Murray) souterraines, Cape Royds, 12 Mars 1908 

Encore la me me " vegetation " problematique, feuilletee, ici d'un blanc sale 
et sans elements figures clairement reconnaissables. Aucuu rhizopode ne s'y est 
rencontre. 

4. Coast Lake, Janvier 1909 

(a) a sec 

(b) sediment dans la formaline 

A part deux exemplaires de Corycia penardi, en mauvais e"tat mais nettement 
reconnaissables encore, et dans lesquels il fut possible de colorer le noyau, le seul 
Rhizopode que parut renfermer ce petit lac a e"te la Quadrula irregularis, Archer, 
une tres petite forme, arrondie, et qui n'e'tait relativement pas tres rare. Elle s'est 
rencontree aussi bien dans le sediment conserve au formol que dans les "herbes" a 
sec, oil, la aussi, on ne la voyait qu'a 1'etat de coquilles vides. 

Mr. Murray avait, du reste, pu examiner sur place ce llhizopode a l'e"tat vivant, 
comme le montre une microphotographie qu'il a bien voulu m'envoyer. 

Cette espece, tres-cosmopolite, aquatique en general mais qui se rencontre egale- 
ment parfois dans les mousses, varie passablement d'une localite a une autre, sous le 
rapport de la compression de 1'enveloppe. Tantot fortement aplatie, tantot presque 
ronde en section transversale, elle se montre ou disco'ide ou globuleuse ; c'est sous 
cette derniere forme, par exemple, qu'on la trouve au fond des lacs suisses (var. globulosa 
PENARD) ; et c'est comme telle egalement qu'on la rencontrait au Cap Rbyds ; beaucoup 
plus petite cependant, la forme de Coast Lake mesurant 23 M, en regard de 30-40 M 
qu'elle atteint dans le lac de Geneve. Au Spitzberg, dans les mousses, on en a trouve" 
une variete tres pen comprime'e egalement. 

C'est dans ce meme Coast-Lake que Mr. Murray a constate la presence d'un 
heliozoaire, Acanthocystis spec., qu'il figure a la PI. VIII. du fascicule " on micro- 
scopic life at Cape Royds," mais que malgre toutes mes recherches je n'ai pas rdussi a 
retrouver. D'apres Murray, cette espece etait abondante en Avril 1908 ; la recolte que 
j'ai examinee e"tait de Janvier 1909. 

5. Clear Lake, Fevrier 1908 

Malgre la presence d'une abondante vegetation microscopique (conferves, algues, 
diatomees minuscules), comme aussi de rotiferes et de quelques infusoires, aucun 
rhizopode ne s'est rencontre dans cette recolte. 



206 E. PENARD 

6. Blue Lake 

(a) ve'ge'tation a sec 

(l>) sediment dans la formaline 

Recolte de J. Murray, en Septembre 1908, a 3 pieds de profondeur, dans la 
partie resserree (narrows) du lac. 

La Quadrula irregular is s'est retrouvee ici, plus abondante meme qu'a Coast 
Lake, et peut-e"tre avec une taille legerement superieure ; d'ailleurs parfaitement 
typique, et, elle aussi, presque ronde en section trail sver sale. 

Dans ce meme petit lac se trouvait Corycia flava (un seul individu), et 
Amoeba terricola. Mais pour cette derniere, je ne puis certifier d'une identi- 
fication absolue ; les deux ou trois organismes qui, d'apres leur apparence generale 
et leur noyau ovo'ide, m'ont paru pouvoir se rapporter a cette espece, etaient 
enkystes, entoures d'une enveloppe speciale, membraneuse, souple et relativement 
epaisse, telle qu'on ne la connait pas dans cette espece, laquelle d'ailleurs, en 
Europe, n'a jamais montrd de phenomenes ve'ritables d'enkystement. 

Probablement est-ce dans ce Blue Lake que Murray a trouve la Nebela vas, 
cette espece si largement repandue dans I' hemisphere austral, et qui parviendrait 
ainsi jusqu'a 1'extreme sud. Mais elle y serait en tout cas tres rare, car il m'a 
ete impossible d'en retrouver le moindre vestige. 

7. Terraced Lake, Cape Barne 
Recolte de J. Murray, 27 Septembre 1908 

Dans les sediments provenant de ce lac, il ne s'est montre, en fait de 
Rhizopodes, qu'une enveloppe vide de Corycia, en trop mauvais etat pour qu'il 
fut possible de la determiner avec certitude, mais qui m'a paru pourtant devoir se 
rapporter a C. penardi. 

Ces memes sediments, pour le dire en passant, etaient rendus interessants par 
la presence de nombreux fragments d'obsidienne, poussieres volcaniques lancees sans 
doute par 1'Erebus ; puis de fibres vegetales, appartenant a des plantes phanerogames 
(meme une fois &, un conifere), et qui sans doute avaient ete apportees de bien loin 
par les vents. 

8. Moraines echouees (stranded moraines) 
Recolte de R. Priestley, Novembre 1908 

Les quelques petits fragments de mousses, recoltes par Mr. Priestley dans les 
" moraines echouees," sur la cote du Continent et a quelque 30 kilometres au sud du 
Cap Royds, ont fourni 1'une des meilleurs recoltes de rhizopodes qui aient e"te faites 
dans 1'Antarctique. 



RHIZOPODES D'EAU DOUCE 207 

Outre la petite Arcelle des mousses, Arcella arenaria, relativement assez com- 
mune, et la Difflugia lucida qui n'etait pas tres rare, on trouvait la un representant 
du genre Diploclilamys, D. timida, nettement caracteristique, 1 puis Euglypha 
compressa, representee, a 1'etat de coques vides, par deux ou trois individus de 
tres faible taille ; enfin Corycia flava. Comme toujours et partout, cette demiere 
espece n'a exige que quelques instants, une fois les mousses humecte'es, pour 
reprendre vie. 

9. Hut Point 
Recolte de R. Priestley, Novembre 1908 

Daus cette recolte, et en depit d'un examen pousse tres a fond, pas un rhizopode 
]ie s'est montre. Les seuls organismes observes ont ete quelques rotiferes enkystes. 
Des spicules de spongiaires, nombreuses, attestaient le voisinage de la mer. 

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Les especes marquees * n'ont etc trouve'es que par Mr. Murray. 

Le tableau qui vient de recapituler les paragraphes precedents, ne sera pas 
sans doute depourvu de quelque intdret, en permettant d'envisager d'un coup d'oeil 
les resultats obtenus. 



1 Tres probablement, la D. fracjilis se trouvait egalement representee ; mais les quelques specimens 
rencontres etaient en si mauvais etat que le fait ne peut etre donn6 cornme certain. 



208 E. PENARD 

Dans cette liste gene'rale, j'ai cru devoir faire entrer, aussi bien que les autres, les 
stations qui se sont montrees vierges de tout Sarcodine. Peut-etre ces colon nes vides 
seront-elles tout aussi significatives, en faisant ressortir, par le fait me'me qu'elles n'ont 
rien a montrer, la pauvrete vraiment extraordinaire des terres antarctiques en rhizo- 
podes d'eau douce. 

Ces parages sont tres pauvres en effet. Peut-etre, au premier coup-d'ceil jete sur 
ces quinze noms d'especes e'numere's les uns au-dessous des autres, sera-t-on tente de 
trouver a redire a cette qualification de " pauvres," de considerer meme le nombre 
obtenu comme un resultat auquel on n'aurait guere pu s'attendre, alors que sous 
les me'mes latitudes on n'avait signale jusqu'ici que trois Rhizopodes, Amoeba 
terricola et Corycia flava, trouves par RICHTERS dans les mousses du Gaussberg, 1 puis 
Arcella arenaria, observes par le meme auteur dans des mousses du Victoria-Land. 2 

Mais ce que la liste ne pouvait pas dire, c'est que cette penurie d'organismes reside 
non pas tant dans le nombre des especes que dans celui des individus. 

Pour qui a quelque experience dans la recherche de ces Sarcodines bryophiles, les 
mousses de 1'Antarctique sont en effet un etonnement. Si Ton excepte la Corycia 
flava, qui dans deux localitcs (Cape Royds et Stranded moraines) se montrait avec 
une proportion d'individus h, peu pres comparable a ce qu'on pourrait s'attendre a 
trouver en Europe, tout est extremement rare. Meme Arcella arenaria, Difflugia 
lucida, que j'indique comme relativement communes, ne se rencontraient que de temps 
ft autre ; et quand aux autres especes, on peut dire, j'en suis sur, que si les precedents 
observateurs ne les ont pas trouvees, c'est qu'il fallait pour cela un specialiste disposant 
de tout son temps, habitue a la patience, et surtout bien decid^ a ne s'attaquer qu'aux 
seuls Rhizopodes pour laisser de cote tout le reste, et a ne considerer Rotiferes, 
Tardigrades, Acariens et Nernatodes que comme des quantites negligeables. 

La plus grande partie, en effet, des especes cities, se sont montrees repre- 
sentees par quelques individus seulement, peniblement obtenus apres de longues 
heures de recherches, et eux-memes en mauvais etat, sous la forme de coquilles 
vides, deformees, et quelquefois brisees. 

On serait meme en droit, semble-t-il, de se demander si ces individus appar- 
tenaient bien a la faune de 1'Antarctique, s'ils n'avaient pas ete apportes la deja 
sans vie, et au meme titre que les fibres de ve'getaux phaneroganiiques qu'on a 
rencontres dans les lacs, par les vents du nord ou par les pattes des oiseaux au 
retour de leurs quartiers d'hiver. 

Mais il ne faut pas nous arreter longtemps a cette idee. Tout en admettant 
que chaque espece ait a 1'origine e'te introduite, et cela dans un passe plus ou 
moins recule, il faut reconnaitre que ces Rhizopodes faisaient, au moment de la 
recolte, partie integraute de la faune locale. L'aspect special des enveloppes, les 
debris que les animaux avaient pu y incorporer, tout cela le montrait deja claire- 



1 Deutsche Siidpol. Exped. 1901-3. Vol. 9. Zool. 

2 Zool. Anzeiger, Juillet 1909, p. 604. 



HHIZOPODES D'EAU DOUCE 209 

ment ; et puis, toute cette faune (a 1'exception de Quadrula irregularis, qui se 
trouve, elle, bien et clument etablie dans les lacs) est une faune spe"ciale, caracte"- 
ristique des mousses, et Ton ne peut concevoir que les vents ou les oiseaux aient 
fait un choix, et n'aient pas su apporter des coquilles ce difflugies, d'arcelles, et 
tant d'autres, qu'ils eussent tout aussi bien pu trouver. 

A mon avis, c'est avant tout dans les conditions climateriques qu'il faut 
chercher Fexplication de cette penurie de Sarcodines ; dans le climat, et aussi, si 
Ton veut, dans les mousses elles-memes. Ces dernieres, en effet, n'arrivent qu'a 
grand peine, dans ces regions du Cap Royds, a trainer leur maigre existence ; 
bien loin d'egaler en vigueur les mousses du Spitzberg, ou Ton peut les recolter 
en touffes splendides, ce ne sont plus ici qui des vegetaux rabougris, dont les 
tiges serrees ne forment qu'un feutre mince et dur ; et dans des conditions 
semblables, au Pole antarctique comme sur les rochers des hautes Alpes, la 
faune microscopique est toujours pauvre. 

Cependant, c'est le climat par lui-meme qui doit ici jouer le role le plus 
direct ; il est decidement trop rude ; s'il ne 1'dtait, il y aurait place encore, dans 
ces mousses si courtes soient-elles, pour une faune rhizopodique bien plus abon- 
dante qu'elle ne Test en re"alite. Dans les lacs, ^galement, si riches en Algues, en 
Rotiferes, Tardigrades, Nematodes et meme Infusoires, on ne trouve a part de temps 
a autre une Corycia qui semble s'etre egaree la que la Quadrula irregularis. 

Dans une lettre qu'il m'ecrivait tout dernierement, Mr. Murray clisait : " Of 
all the moss-dwellers, I find the Rhizopods least willing to come alive after being 
for a long time dried." La conclusion du biologiste du Nimrod est celle-la meme 
a laquelle mes etudes m'avaient depuis longtemps amend, et que les recoltes de la 
derniere expedition antarctique vienuent une fois de plus confirmer. Partout les 
memes faits se repetent: Acariens, Ndmatodes, Tardigrades et Rotiferes recom- 
mencent a courir alors que les Rhizopodes gardent encore un repos absolu, ou 
reprennent vie alors que ces derniers ne le pourront jamais plus. 

Les Rhizopodes, d'une maniere generale, sont moins resistants que les orga- 
nismes qui vienuent d'etre cites ; mais il ne faudrait pas envisager cette donnee 
comme absolument generale. Parmi les Sarcodines des mousses, il en est quel- 
ques-uns qui constituent une exception bien nette a la regie ; je citerai par 
exemple Diffluyia arcula, puis Bullinula indica, qui rivalisent sous ce rapport 
avec les Kotiferes les plus endurants (et pourtant manquaient au Cap Royds, pour 
des raisons speciales sans doute) ; puis surtout la Corycie, Corycia flava, qui, elle, 
est d'une resistance incroyable, et dans cet ordre de faits tient peut-etre le record 
dans toute la serie des organismes animaux. 

En resume, les collections rapportees de 1'Antarctique, et qui ont donne a 
Mr. J. Murray de si interessants re"sultats pour les groupes qu'il a etudies, en ont 
fourni de quelque valeur egalement pour les Sarcodiues. Elles ont montrd que ces 
organismes se trouvent, en fait, representes la-bas par les memes especes que 



210 E. PENARI) 

partout ailleurs; que ces especes y sont, mais representees par uue tres petite 
quantite d'individus. 

Toute cette faune est arrivee, sans doute, tout pres de la limite au-dela de 
laquelle la vie lui reste ferme'e au sud ; mais cette limite, 1'homme 1'a depassee, 
Shakleton et ses compagnons, pour aller planter a deux pas du Pole le pavilion du 
Royaume Uni. 



CHAPITRE II 

COLLECTIONS RECUEILLIES PENDANT LE VOYAGE DE RETOUE 

A. ILES DU PACIFIQUE SUD 

1. lie Macquarie 

Recolte effectuee par Capt. J. K. Davis a Lusitania Bay, 
pres d'un marecage. Mai 1909 

(a) Une touffe d'une mousse longue et serree, et un paquet de conferves. 

(b) Sediment dans la formaline. 



Amoeba terricola. 
Arcella arenaria. 
vulgaris. 
Assulina inuscorum. 
Bullinula indica. 
Corythion dubium. 
Difflugia arcula. 

constricta. 
Euglypha alveolata. 
ciliata. 
compressa. 
laevis. 



Heleopera petricola. 

sphagni. 
sylvatica. 
Nebela caudata. 
dentistoma. 
lageniformis. 
martiali. 

vas. 

Phryganella hemisphaerica. 
Sphenoderia dentata. 
Trinema enchelys. 



2. lies Auckland 

Fragment de mousse re'colte'e par le Dr. Cockayne, botaniste du 
Gouvernement de la N 1Ie Zelande. 1907 

Cette mousse tapissait le tronc d'un "Eata," Metrosideros lucida 



Assulina muscorum. 
Difflugia arcula. 

constricta. 
BRIT. ANTARCT. EXPED. 1907-9 VOL. I. 



Euglypha compressa. 
Hyalosphenia cockayni. 
Nebela tincta. 



B 



E. PENARD 



B. NOUVELLE ZELANDE 

1. lie Stewart 
Re"colte de J. Murray, au sommet d'un petit ilot. Mars 1909 



Amoeba terricola. 
Arcella arenaria. 
Assulina muscorum. 
seminuluni. 
Bullinula indica. 
Centropyxis horrida. 
Corythion dubium. 
Difflugia arcula. 

constricta. 



(a) Mousses seches, en petite quantity. 
(6) Sediment dans la formaline. 

Diplochlamys fragilis. 
Euglypha ciliata. 

compressa. 

laevis. 
Heleopera sylvatica. 
Nebela lageniformis. 
Phryganella hemisphaerica. 
Plagiopyxis callida. 
Trinema enchelys. 



2. District du Mont Cook, He du Sud 

* Recoltes de J. Murray, sur diffe'rents points, de 3000 a 6000 pieds d'altitude, 
et sur un espace de plusieurs milles. Dccembre 1907 

(a) Mousses diverses, avec une belle touffe de sphagnum. 
(6) Sediment dans la formaline. 



Amoeba sphaeronucleolus. 

terricola. 
Arnphizonella violacea. 
Arcella arenaria. 
Assulina muscorum. 

seminulum. 
Bullinula indica. 
Centropyxis aculeata. 
Corycia penardi. 
Corythion dubium. 
Cyphoderia ampulla. 
Difflugia arcula. 

constricta. 

lucida. 

manicata. 

pyriformis, var. bryophila. 
Diplochlamys timida. 
Euglypha alveolata. 
ciliata. 



Euglypha compressa. 
Heleopera petricola. 
sphagni. 

Hyalosphenia cockayni. 
Nebela collaris. 
dentistoma. 
lageniformis. 
longicollis. 
tincta. 

vas. 
Phryganella hemisphaerica. 

nidulus. 
Pontigulasia bryophila. 
compressa. 

Sphenoderia dentata. 

fissirostris. 

Trinema enchelys. 
lineare. 



RHIZOPODES D'EAU DOUCE 



213 



**Re"colte du Dr. Mackay, stir un seul point, a Nun's Veil, 
5000 pieds. Decembre 1907 



Arcella arenaria. 
Bullinula indica. 
Difflugia arcula. 

constricta. 
Euglypha ciliata. 



Euglypha laevis. 
Nebela lageniformis. 

vas. 
Phryganella hemisphaerica. 



3. Waiata-Eua, Cote occidentals, He du Nord 

Recolte de J. Murray, dans le "bush," pres des chutes -du Nihotupo. 
Mousses diverses, en petite quantite. Decembre 1907 



Amoeba terricola. 
Assulina muscorum. 

seminulum. 
Bullinula indica. 
Corythion dubium. 
Difflugia arcula. 

constricta. 

lucida. 
Diplochlamys fragilis. 
Euglypha ciliata. 

compressa. 



Euglypha strigosa. 
Heleopera sphagni. 
Nebela collaris. 

,, lageniformis. 

longicollis. 

tincta. 

vas. 

Phryganella hemisphaerica. 
Plagiopyxis callida. 
Pontigulasia compressa. 
Trinema enchelys. 



4. Ngauruhoe, volcan, He du Nord 

Recolte du Dr. Mawson, dans une seule region, a 4-5000 pieds 

d'altitude. Avril 1909 



(a.) Mousses diverses, a sec. 
(6) Sediment dans la formaline. 



Assulina rnuscorum. 

seminulum. 
Bullinula indica. 
Centropyxis laevigata. 
Corythion dubium. 
Difflugia arcula. 

constricta. 

lucida. 
Euglypha ciliata. 

,, compressa. 



Heleopera sphagni. 
Nebela caudata. 

collaris. 

lageniformis, 

martiali. 

militaris. 

vas. 

Phryganella hemisphaerica. 
Trinema enchelys. 



214 



E. PENARD 



5. Rotorua, He du Nord 
Recolte de J. Murray, dans le "bush." Avril 1909. Mousses diverses 



Amoeba terricola. 
Arcella arenaria. 
Bullinula indica. 
Centropyxis laevigata. 



Difflugia constricta. 
Euglypha ciliata. 
Nebcla lageniformis. 
Phryganella hemisphaerica. 



C. AUSTRALIB 



1. Sydney, Nouvelle Galles du Sud 



Mousses diverses, recoltees par J. 
puis dans les environs de 



Amoeba muralis. 

radiosa. 

terricola. 
Assulina muscoruin. 

seminulum. 
Bullinula indica. 
Corycia flava. 
Corythion dubium. 
Difflugia arcula. 

constricta. 

fallax. 
Euglypha ciliata. 

compressa. 

laevis. 
Heleopera petricola. 



Murray, au National Park, 
Sydney. Avril 1909 

Heleopera sordida. 

sylvatica. 

Hyalosphenia cockayni. 
Nebela certesi. 
collaris. 
lageniformis. 
martiali. 
tincta. 

vas. 

Parmulina cyathus. 
Phryganella hemisphaerica. 
Plagiopyxis callida. 
labiata. 

Trinema complanatum. 
enchelys. 



2. Alpes australiennes, Nouvelle Galles du Sud 
Recoltes de J. Murray. Avril 1909 

(a) Region tourbeuse pros de 1'Hospice, 5000 pieds ; mousses et sphagnum. 
(6) "The Creel," 3000 pieds, mousses, belles et longues. 
(c) Sddiment dans la formaline. 



Amoeba terricola. 
Amphizonella violacea. 
Arcella arenaria. 



Assulina muscoruin. 
Bullinula indica. 
Corycia flava. 



RHIZOPODES D'EAU DOUCE 



215 



Corythion dubium. 
Difflugia arcula. 

constricta. 

fallax. 
Diplochlamys fragilis. 
Euglypha alveolata. 

ciliata. 

compressa. 

filifera. 

laevis. 

strigosa. 
Heleopera petricola. 
sylvatica. 



Lieberkiihnia wageneri. 
Nebela caudata. 
lageniformis. 
martial! . 

vas. 

Phryganella hemisphaerica. 
Plagiopyxis callida. 
labiata. 

Quadrula symmetrica. 
Sphenoderia dentata. 
Trinema complanatum. 
lineare. 



3. Katoomba, Montagnes bleues, N llc Galles du Sud 

llecoltes de J. Murray, dans trois petites gorges (gullies) qui menent a la 
"Sunken Valley," et sur un territoire de 2 milles. Mai 1909 

(a) Mousses diverses, avec une touffe de sphagnum. 
(&) Sediment dans la formaline. 



Amoeba sphaeronucleolus. 

terricola. 
Amphitrema stenostoma. 
Amphizonella violacea. 
Arcella arenaria. 
Assulina muscorum. 

seminulum. 
Bullinula indica. 
Centropyxis horrida. 

laevigata. 

Corycia flava. 
Corythion dubium. 
Difflugia arcula. 

constricta. 

lucida. 
Diplochlamys fragilis. 
vestita. 

Euglypha compressa. 



Euglypha cristata. 
laevis. 
strigosa. 
Heleopera sylvatica. 
Nebela certesi. 
collaris. 
griseola. 
lageniformis. 
longicollis. 
martiali. 
militaris. 
tincta. 

vas. 

Phryganella hemisphaerica. 
Plagiopyxis callida. 
Trinema complanatum. 
enchelys. 
lineare. 



216 



E. PENARD 



4. Eumundi Station, Queensland 

Recolte de J. Murray, dans le " bush," sur un territoire de deux milles 

cane's environ. Mai 1909 

(a) Paquet de mousses, sans sphagnum. 

(&) Beau paquet de mousses varie"es, quelquefois sur ecorce, avec un petit fragment de 
sphagnum. 

(c) Sediment dans la formaline. 



Amoeba sphaeronucleolus. 

terricola. 
Arcella arenaria. 
Assulina muscorum. 
Bullinula indica. 
Centropyxis aculeata. 

delicatula. 

Corycia flava. 
Corythion dubium. 
Difflugia arcula. 

constricta. 

lucida. 

pyriformis, var. bryophila. 



Diplochlamys timida. 
Euglypha ciliata. 
compressa. 
laevis. 
Heleopera sphagni. 
Hyalosphenia subflava. 
Nebela caudata. 

collaris. 

,, lageniformis. 
Phryganella hemisphaerica. 
Plagiopyxis callida. 
Trinema complanatum. 

enchelys. 



D. ILES DU PACIFIQUE 
1. Suva, Fiji 

K,e"colte de J. Murray, pres du rivage, au bord d'un marecage a paletuviers, puis 

dans 1'interieur, a un mille ou deux de la cote, dans les fosses 

au bord de la route. Mai 1909 

(a) Mousses tres-courtes, avec beaucoup de terre. 

(b) Sediment dans la formaline. 



Amoeba terricola. 
Bullinula indica. 
Centropyxis delicatula. 
Corycia aculeata. 

penardi. 
Difflugia arcula. 

constricta. 
lucida. 



Difflugia pyriformis, var. bryophila. 

Euglypha laevis. 

Heleopera petricola. 

Nebela vas. 

Phryganella hemisphaerica. 

Quadrula irregularis. 

Trinema lineare. 



RHIZOPODES D'EAU DOUCE 



217 



2. Oahu, Hawaii 

Recolte de J. Murray, aux environs d'Honolulu, dans une vallde ou le " bush " 
devint rapidement impraticable a cause de la densite de la vegetation, les 
fougeres (Pteris) couvrant si bien le sol que la marche s'efFectuait h, un ou 
deux yards au-dessus du sol. Mai 1909 

(a) Belles mousses, 1'une d'elles tres-claire, longue et soyeuse. 

(b) Sediment dans la formaline. 



Amoeba terricola. 
Assulina seminulum. 
Bullinula indica. 
Centropyxis aculeata. 
Corycia flava. 
Corythion dubium. 
Cyphoderia ampulla. 
Difflugia arcula. 

constricta. 
Diplochlamys gruberi. 
timida. 



Euglypba ciliata. 
,, compressa. 
laevis. 
Heleopera sylvatica. 
Nebela collaris. 

lageniformis. 

militaris. 

minor. 

Phryganella hemisphaerica. 
Sphenoderia dentata. 
Trinema enchelys. 



E. COLOMBIE BEITANNIQUE ET CANADA 



1. Victoria, B.C. 

Rdcolte de J. Murray, parmi les plantes du rivage (Sedum, &c.), puis 
dans un petit bois, non loin du rivage dgalement. Juin 1909 



Amoeba terricola. 
Arcella arenaria. 
Assulina muscorum. 
Bullinula indica. 
Corytbion dubium. 
Difflugia constricta. 
Euglypha ciliata. 



Euglypba laevis. 

strigosa. 
Nebela bigibbosa. 

caudata. 

Phryganella hemispbaerica. 
Plagiopyxis labiata. 
Trinema enchelys. 



218 



E. PENARD 



1*. Vancouver, B.C. 
Recolte de J. Murray, Stanley Park. Juin 1909 

(a) Mousses, avec une touffe de sphagnum. 
(1) Sediment dans la formaline. 



Arcella arenaria. 
Assulina muscorum. 

,, seminulum. 
Centropyxis aculeata. 
Corycia flava. 

penardi. 
Cyphoderia ampulla. 
Difflugia arcula. 

bacillifera. 

constricta. 

lucida. 

pyriforrnis, var. bryophila. 
Diplochlamys vestita. 
Euglypha alveolata. 

brachiata. 

ciliata. 

compressa. 

strigosa. 
Heleopera petricola. 
sordida. 



Lesquereusia 

Lieberkiihnia wageneri. 
Nebela bigibbosa. 

caudata. 
collaris. 
dentistoma. 
flabelluliiii). 
lageniformis. 
tincta. 
tubulosa. 

vas. 

Phryganella hemisphaerica. 
Placocysta jurassica. 

spinosa. 
Plagiopyxis callida. 
Pontigulasia bryophila. 
Quadrula symmetrica. 
Trinema complanatum. 
lineare. 



2. Monts Selkirk, B.C., et Montagues Rocheuses, Canada 

Recoltes de J. Murray, a des altitudes de 3000 a 6000 pieds, aupres des diverses 
stations de la ligne ou le train s'arretait plus de 5 minutes de temps. Juin 1909 

(a) Mousses diverses. 

(&) Sediment dans la formaline. 



Amoeba sphaeronucleolus. 

terricola. 
Arcella arenaria. 
Bullinula indica. 
Corycia flava. 
Difllugia arcula. 

constricta. 



Euglypha ciliata. 

compressa. 
Nebela collaris. 

lageniformis. 
Phryganella hemisphaerica. 
Plagiopyxis callida. 



RHIZOPODES D'EAU DOUCE 



219 



3. Province d' Ontario, Canada 
+ Lake of the W^oods 

Re"colte de J. Murray, dans le fourre, au bord du lac, pendant 
un arr6t du train. Juin 1909 



(a) Mousses diverses. 

(&) Sediment dans la formaline. 



Amoeba sphaeronucleolus. 

terricola. 
Arcella arenaria. 
Assulina muscoruin. 

seminulum. 
Bullinula indica. 
Corythion dubium. 
Difflugia constricta. 



Euglypha ciliata. 
compressa. 
laevis. 
Nebela lageniformis. 

tincta. 

Phryganella hemisphaerica. 
Trinema enchelys. 
lineare. 



H 



Ottawa 



Recolte de J. Murray, dans le " Public Park," et tout aupres 
sur le bord du chemin. Juin 1909 

(a) Mousses seches. 

(b) Sediment dans la formaline. 



Amoeba terricola. 
Arcella arenaria. 
Awerinzewia cyclostoma. 
Corycia flava. 

penardi. 
Difflugia constricta, 

lucida. 
Diplochlamys fragilis. 



Diplochlamys timida. 
Euglypha ciliata. 
laevis. 
Nebela minor. 
Parmulina cyathus. 
Phryganella hemisphaerica. 
Trinema complanatum. 
lineare. 



BEIT. ANTARCT. EXPED. 1907-9. VOL, I. 



E. PENARD 



RECAPITULATION 





ANTARCTIQUE 


PACI- 

FIQUE 
SUD 


N LL " ZELANDE 


AUSTRALIE 


PACI- 

FIQUE 


COLOMBIE 
ET 

CANADA 


ILE MACQUARIE 


ILES AUCKLANDS 


ILE STEWART 


T3 

02 

3 
T3 
IB 

t I 

M 

o 
o 

a 

f4 

i~^ 

<! 


Tg 

o 
K 

s 
n 

a) 
Fn 

4 
& 

M 
4 

H 
j 

5 
f 


NGAURUHOE, He du Nord 


"2 

o 

& 

a 
~a 



KH 

f 
& 



i 

o 

M 


K 

E 

a 
f 

cc 


ALPES AUSTRALIENNES 


KATOOMBA 


EUMUNDI, Queensland 


S 

< 

> 

b 

02 


1 

W 

tT 
a 
t 
O 


VICTORIA ET VANCOUVER 


M TS SELKIRK, M KS ROCHEUSES 


ONTARIO 


Amoeba muralis .... 






























































sphaeronucleolus . 
terricola .... 
Amphitrema stenostoma 
Ainphizonella violacea 
Arcella arenaria .... 
vulgaris ..... 
Assuliua muscorum .... 
seminulum .... 
Aweriuzewia cyelostoma 
Bullinula indica .... 
Centropyxis aculeata .... 
,, delicatula 
,, horrida .... 
laevigata 
Corycia aculeata .... 
,, flava ..... 
penardi ..... 
Corythion dubiura .... 
Cyphoderia ampulla .... 
Difflugia areula ..... 
,, bacillifera .... 
,, constricta .... 
fallax ..... 
,, lucida .... 
,, mauicata .... 
,, pyriformis, var. bryophila 
Diplochlamys fragilis .... 
gruberi . 
,, timida .... 
,, vestita .... 
Euglypha alveolata .... 
brachiata .... 
ciliata .... 
,, compressa .... 
,, ciistata .... 
filifera .... 
,, laevis ..... 
strigosa .... 


1 


4- 


4- 


+ 
+ 


+ 





4- 


4- 


4- 


4- 
4- 


4- 
4- 


+ 


+ 


+ 


4- 
4- 


4- 
4- 


+ 


+ 

+ 
+ 


4- 


+ 
+ 








4- 





4- 

4- 


4- 

4- 


+ 








4- 


4- 


4- 


+ 


+ 


+ 


+ 

+ 


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4- 


4- 
4- 





+ 
4- 


4- 


4- 
4- 


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+ 


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4- 





4- 
4- 
4- 

4- 


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+ 


+ 

| 


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+ 


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4- 


4- 


4- 


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4- 


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4- 
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4- 


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4- 


4- 
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' _j_ 


4- 








4- 
















4- 

+ 


+ 





4- 


-t- 
+ 


+ 


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4- 
4- 


4- 

+ 


4- 
4- 


4- 
4- 


4- 


4- 

4- 
4- 
4- 


4- 

4- 
+ 
4- 
4- 


4- 

4- 


4- 
4- 

4- 





4- 


4- 


4- 


+ 


4- 


4- 





4- 


4- 


4- 


4- 


4- 


+ 


+ 


+ 


4- 


+ 
4- 


4- 

4- 


4- 

4- 


4- 


4- 

4- 


4- 
4- 


4- 

4- 


4- 

4- 


4- . 
4- 


4- 


f 

4- 


4- 


4- 
4- 











4- 


+ 

4- 











4- 


4- 


4- + 




4- 





4- 


+ 




























4- 




















4, 













+ 


























































4- 


4- 
+ 


4- 


4- 

+ 


4- 

4- 


4- 
4- 


4- 
4- 


4- 


4- 

4- 


4- 

4- 


4- 
4- 


4- 

4- 


4- 
4- 


4- 
4- 


4- 
4- 


4- 
4- 
































4- 


+ 





+ 


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4- 


4- 

4- 


4- 
4- 


+ 4- 


4- 


4- 

4- 


i 

i 



RHIZOPODES D'EAU DOUCE 



RECAPITULATION (suite) 





ANTARCTIQUE 


PACI- 

FIQUE 
SUD 


N MJ! ZELANDE 


AUSTKALIE 


PACI- 

FIQUE 


OOLOMBIE 

ET 

CANADA 


ILE MACQUARIE 


ILES AUCKLANDS 


ILE STEWART 


T3 

EC 

TS 
,2 
H 

M" 

8 
o 

H 
3 


3 
& 

o 

9 

H 
f 

& 

P5 
-i 

H 

jj 

^ 


NGAURUHOE, He du Nord 


13 
t-t 


K 

ff 
T3 

a 

HH 
< 

I 

O 
H 
O 

PH 


SYDNEY 


ALPES AUSTHALIENNES 


->! 


S3 

<t 

M 


EUMUNDI, Queensland 


5? 

4 

> 

o 

m 


: 3 

r 

<s 

w 

cT 
i 

<i 



VICTORIA ET VANCOUVEK 


M TS SELKIRK, M KS 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', B d 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., B d 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., B a 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., B d 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., B d 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., B d 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., B d 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, M es 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 2 e 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., B d 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 Isl d , 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) Cdte 1 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 P RIC E Is. 6d. NET 

VOL. I. PART II., ON MICROSCOPIC LIFE AT CAPE ROYDS 

Bv JAMES MURRAY P RIC E 5s. Od. NET 

VOL. I. PART III., ANTARCTIC ROTIFERA 

BY JAMES MURRAY P R ,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 P R ,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 





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. 

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 f i. 
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. 7 C 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<?) 

Ph. filis non in strato associatis, sed inter algas varias Myxophycearum libere 
natantibus, brevibus ; vaginis indistinctis, plerumque diffluentibus ; trichomatibus 
brevibus vel brevissimis, pallide serugineis, valde et subirregulariter spiraliter contortis, 
apicibus obtusis et non attenuatis; cellulis diametro trichomatis 1-2-plo longioribus, 
contentu homogeneo. 

Crass, trich. 0.6 M; long. cell. 0.6-1.2 M. 

Hob. Pony Lake. 

This minute and much- twisted Phormidium was present in quantity in the 
sediment obtained from under the ice of Pony Lake both in April 1908 and in 
January 1909. It was not observed in any of the other collections, and occurred 
mixed with various Green and Blue-green Algse. The trichomes are very short 
and invariably twisted into some form of spiral, although they exhibit great 
irregularity. 

It need not be confused with any of the spirally twisted piankton species 
of Lyngbya. It has no definite sheath, and is smaller and narrower than any of 
them. Some specimens showed slight indications of a sheath which had become 
diffluent, otherwise the Alga might be regarded as an irregular species of the genus 
Arthrospira. 



FRESHWATER ALGM 293 

Genus OSCILLATORIA Vaucher 

62 Oscillatoria limosa Ag. 

1812; Gomont, I.e. p. 230, t. 6, f. 13. 

Crass, trich. 14-17 n. 

Hob. Pony Lake, Coast Lake, and Green Lake. Also ponds on Mt. Erebus. 

This species was abundant in the above-mentioned localities, and in most cases 
was remarkable for the shortness of its filaments, many of which were only 40-60 M in 
length. 

63 Oscillatoria sancta Kiitz. 

Tab. Phycol. I, p. 30, t. 42, f. 7 ; Spec. Alg. 1849, p. 246 ; Gomont, l.c. p. 229, 
t. 6, f. 12. 

Crass, trich. 12.5-15 M. 
Hob. Pond, Cape Barne. 

64 Oscillatoria subproboscidea sp. n. (PI. XXV, Figs. 91-94) 

0. trichomatibus inter varias Oscillatorias libere natantibus, olivaceo-eerugineis, 
elongatis, rectis vel leviter flexuosis, juxta apicem attenuatis et plerumque subuncinatis, 
inter cellulas non constrictis ; cellulis diametro trichomatis circiter 2|-plo brevioribus, 
non elongatis apicem versus; cytioplasmate minutissime granulatis, dissepimenta 
non granulata; cellula apicali brevi et convexo-obtusa, membranam convexam leviter 
incrassatam praebens. 

Crass, trich. 8.2-9 M; long. cell. 3-4 ft. 

Hab. Coast Lake. 

This species was frequent in the sediment under the ice of Coast Lake, mixed with 
0. tennis and others. It is perhaps nearest to 0. proboscidea Gomont, but is consider- 
ably narrower, with straight trichomes and rather longer cells. The apex is also 
different, as although it is somewhat suddenly attenuated and frequently uncinate, 
it is never capitate. 

In thickness 0. subproboscidea is similar to 0. chalybea, but there are no constric- 
tions between the cells as in the latter species, and the trichomes are only attenuated 
close to the apex. The apical cell is also short and possesses a calyptra. 

65 Oscillatoria tennis Ag. 

1813 ; Gomont, l.c. p. 240, t. 7, f. 2, 3. 

Crass, trich. 7-10 n. 

Hab. Coast Lake. 

This species was frequent amongst 0. subproboscidea and a small quantity of 
0. limosa Ag. The trichomes were typical in all respects, and varied considerably 
in thickness. 



294 W. AND G. S. WEST 

66 Oscillatoria producta sp. n. (PI. XXV, Figs. 86-90) 

0. trichomatibus sparsis inter alias algas Myxophycearum natantibus, serugineis, 
subrectis, inter cellulas leviter constrictis, apices versus et juxta leviter attenuatis ; 
cellulis diametro trichomatis 3-4- plo brevioribus; cytioplasmate granulata, dissepi- 
menta non granulata ; cellula apicali in processum mamillatum vel digitatum 
nonnunquam uncinatum, p^us minusve producta, usque diametro 2|-plo longiori, 
pellucida, ssepe calyptra convexo-conica ornata. 

Crass, trich. 5.3-6.8 n ; long. cell. 1.4-1.8 n. 

Hob. Pond on Mt. Erebus. 

Perhaps the nearest species to 0. producta is the one described as 0. ttoydiana by 
Gomont (in Bull, Soc. botan. France, xlvi, 1899. p. 39, t. 1, f. 17), from which it is 
distinguished by its narrower trichomes, the absence of granules at the dissepiments, 
the differently attenuated apex, and the presence of a calyptra. 

It might also be compared with 0. janthophora Gomont, 0. cortiana Menegh., and 
0. oJceni Ag , from all of which it is easily distinguished. 

67 Oscillatoria cortiana Menegh. 

1837 ; Gomont in Ann. Sci. Nat. T serie, Bot. xvi, 1892, p. 251, t. 7, f. 17.. 
Forma cellulis paullo brevioribus; crass, trich. 5.8-6.9 /* ; long. cell. 4-5.5 ju; 
crass, apic. trich. 2.8-3.3 /u ; long. cell. apic. 7.5-10 ju. 

Hob. Clear Lake, rather scarce among the sheets of Phormidium glaciale. 

68 Oscillatoria priestleyi sp. n. (PI. XXVI, Figs. 99-101) 

0. trichomatibus libere natantibus, densissime aggregatis, pallide serugineis, 
elongatis et subrectis, apicem versus longe attenuatis, apicibus curvatis et s^pe 
recurvatis, interdum irregulariter flexuosis, obtusis, inter cellulas non constrictis, 
hinc inde massa intercellular! inflate- tor ulosa et refringenti interrupts; cellulis 
subquadratis vel plerumque paullo brevioribus, interdum paullo tumidulis, non 
elongatis apicem versus ; cytioplasmate minutissime granulatis et interdum cum 
granulo magno subrefringenti ornato ; cellula apicali obtusa, nunquam capitata ; 
calyptra nulla. 

Crass, trich. 5-5.9 M ; crass, apic. trich. 2.3-3.2 /x; long. cell. 3-5 n. 

Hob. Lake on west side of McMurdo Sound. 

The only species related to Oscillatoria priestleyi are 0. subitliformis Kiitz., 
0. cortiana Menegh., and 0. tanganyikce G. S. West, with the first of which it should 
be very carefully compared. 

0. priestleyi differs from 0. subuliformis in the entire absence of undulation from 
the straight or gently flexuose filaments, in the much more irregular curvature of 
their attenuated apices, in the relatively shorter cells, which remain short to the very 
apex of the filament, and in the presence of numerous refractive lens- shaped masses 
of deeply coloured and deeply staining intercellular substance. The trichomes are 



FRESHWATER ALGJS 295 

of the same thickness as the marine 0. subuliformis, and the gradual attenuation 
towards the apex is a very similar character. 

69 Oscillatoria formosa Bory 

1827 ; Kutz. Tab. Phycol I, t. 41, f. 8 ; Gomont, I.e. p. 250, t. 7, f. 16. 

Crass, trich. 4.8 p. 

Hob. Coast Lake, scarce among 0. tennis, 0. subproboscidea, and 0. limosa. 

70 Oscillatoria chlorina Kutz. 

Phyc. gener. 1853, p. 185 ; Spec. Alg. 1849, p. 239 ; Tab. Phycol. I, p. 28, t. 39, 
f. iii; Gomont, l.c. p. 243. 

Crass, trich. 3.5-4 M; long. cell. 3.8-5.5 M. 
Hob. Green Lake. In pond, Cape Koyds. 

71 Oscillatoria terebriformis Ag. 

1827 ; Gomont, l.c. p. 254, t. 7. f. 24. 
Forma tennis. (PI. XXVI, Fig. 98) 

Forma trichomatibus 3 M crassis, cellulis subquadratis ; dissepimenta granulata; 
cellula apicali subtruncata. 

Hob. In pond, Cape Royds. 

72 Oscillatoria amphibia Ag. 

1827 ; Gomont, l.c. p. 241, t. 7, f. 4, 5. 
Crass, trich. 2-2.1 n. 

Hab. Coast Lake, infrequent among various Myxophyceae. 
Var. robusta var. n. (PI. XXVI, Fig. 102) 

Var. trichomatibus crassioribus ; dissepimenta granules plures pulcherrime 
coaruleas utrobique prsebens. 
Crass, trich. 3.5 M. 
Hob. In pond, Cape Royds. 

73 Oscillatoria deflexa sp. n. (PL XXV, Figs. 95-97) 

0. trichomatibus sparsis et nonnunquam in csespitibus parvis subspiraliter contortis 
inter alias Myxophycearum natantibus, rectis, serugineis, apices versus gradatim 
attenuatis et juxta apicem subiter deflexis; inter cellulas non constriotis, dissepi- 
menta non granulata ; cellulis diametro trichomatis 2-2^-plo longioribus, cytioplasmate 
homogeneo ; cellula apicali attenuata obtusa, calyptra nulla. 

Crass, trich. 0.9-1 n ; long. cell. 2.4-2.9 M ; long. part, deflex. apic. 4.2-4.8 /u. 

Hab. Green Lake. Coast Lake. 

This species should be compared with 0. subtilissima Kiitz., 0. Kutzingiana Nag., 
and 0. angustissima W. & G. S. West, from all of which it differs in the apex of the 
trichomes and other points. 0. minuta Hansg. may also be an allied species. 

BRIT. ANTARCT. EXPED. 1907-9. VOL. I. N 



296 W. AND G. S. WEST 

CHROOCOCCACE^E 
Genus ASTEBOCYSTIS Gobi 

74 Asterocystis antarctica sp. n. (PI. XXVI, Figs. 103-106) 

A. minuta, inter varias algas libere natantes ; filis brevibus, vaginatis, vix ramosis, 
valde contortis, e serie singula cellularum formatis ; vaginis arctis, achrois et 
gelatinosis; cellulis ante divisionem ellipsoideis, post divisionem subhemisphsericis 
vel elongato-hemisphsericis, tarn longis quam latis vel brevioribus ; cytioplasmate 
a3rugineo et granuloso (verisimiliter ut in Chroococco minuto). 

Long. cell. 4.5-8 M ; lat. 6-8 /" ; crass vag. 9-10 /". 

Hob. Green Lake. 

Asterocystis antarctica is of about the same size asA.africana G. S. West (inJourn. 
Linn. Soc, bot. xxxviii, 1907, p. 196), but is of a different habit and has cells of a 
different form. The short, contorted filaments, almost without any trace of branches, 
distinguish it at once from all the other species of the genus. 

Genus MICROCYSTIS Kiitz. 
75 Microcystis chroococcoidea sp. n. (PL XXVI, Figs. 107-114) 

M. coloniis parvis, libere natantibus, e cellulis paucis (6-24) compositis, tegumento 
gelatinoso et achroo, vix conspicuo ; cellulis confertis sed irregulariter aggregatis, iis 
periphericis interdum paullo minoribus, contentu pallide flavo-a3ruginoso, subtiliter 
granuloso cum granulis conspicuis majoribus 2-4. 

Diam. cell. 4-7 /" ; diam. colon. 14-33 n. 

Hob. Green Lake. 

This member of the Chroococcacese was not uncommon in the strongly saline 
water of Green Lake. The colonies are very small, free-floating, and often do not 
consist of more than a dozen cells. The cells are closely aggregated to form a some- 
what irregular colony, in which the peripheral cells are sometimes slightly smaller 
than the others. The cells do not possess an independent envelope, but are all enclosed 
in a very thin and almost invisible mucous integument. The alga forms a connecting 
link between the genera Microcystis and Chroococcus, and might equally well be 
named " Chroococcus microcystoidea," 

Genus GLCEOCAPSA Kiitz. 
76 Glosocapsa. shuttleworthiana Kiitz. 

Phyc. gener. 1843, p. 175 ; Tab. Phycol. I, p. 18, t. 23, f. 1 ; Forti, Syll. 
Myxophy. 1907, p. 37. 

Diam. cell. 2.5-3.5 n, c. integ. prim. 7-8 M ; tegumentis internis intense aurantio-rubris. 

Hob. On stones, High Moraines, Mt. Erebus. Also fragmentary in pond, Cape 
Royds. 



FRESHWATER ALGM 297 

Genus APHANOCAPSA Nag. 

77 Aphanocapsa montana Cramer 

Rabenh. Flor. Europ, Alg. II, 1865, p. 50 ; Forti, Syll. Myxophy. 1907, p. 72. 

Diam. cell. 2-2.4 M. 

Hob. On stones, High Moraines, Mt. Erebus. 

The cells were fairly crowded, and formed yellow- green, gelatinous colonies from 
800 to 1300 /x in extent, closely adherent to the stones, and amongst the thalli of 
Prasiola antarcticu Kiitz. 

Genus CHROOCOCCUS Nag. 

78 Chroococcus minutus (Kiitz.) Nag. 

Gatt. einzett. Alg. 1849, p. 46; Forti, Syll. Myxophy. 1907, p. 14. 
Diam. cell. 8-11 M; colon, usque ad 60 M diam. 
Hab. Green Lake. Pond on Mt. Erebus. 

Var. obliteratus (Richter) Hansg. Prodr. Algenfl. Bohm. II. p. 162. Chr. obliteratm 
Richter in Hauck & Richter, Phyk. Univ. No. 41 ; Notarisia, 1886, p. 97. 
Diam. cell. 6-8 M; diam. colon. 23-29 M. 
Hab. Coast Lake. 

79 Chroococcus minor (Kiitz.) Nag. 

Gatt. einzell. Alg. 1849, p. 47, t. 1A ; Rabenh. Flor. Europ. Alg. II, 1865, p. 30 

Diam. cell. 4 M ; diam. colon. 18-20 M . 

Hab. In pond, Cape Royds. 

Forma minitna West in Journ. Linn. Soc. bot. 1894, p. 275, t. 16, f. 18. Chr. 
minutus var. minimus v. Keissler in Verhandl. der zool.-bot. Ges. Wein, 1901, p. 394, 
f. 1, 2. Chr. minimus (v. Keissler) Lemm. in Archiv. fur Botan. utgifv. af K. Sv. Vet.- 
Akad. Bd. 2, No. 2, 1904, p. 102. 

Diam. cell. 1.9-2.2 M ; diam. colon. 22-31 /u. 

Hab. Green Lake. Lake on west side of McMurdo Sound. 

80 Chroococcus pallidus Nag. 

Gatt. einzell '. Alg. 1849, p. 46, t. 1, f. A2 ; Rabenh. Flor. Europ. Alg. II, 1865, p. 33. 
Diam. cell. 6-7 M; diam. colon. 16-21 M. 
Hab. Lake on west side of McMurdo Sound. 

The colonies were rather irregular and consisted of four or eight almost globose 
cells. The cell- contents were pale yellow- green and granulose. 

81 Chroococcus cohcerens (Breb.) Nag. 

Gatt. einzell. Alg. 1849, p. 46 ; Rabenh. Flor. Europ. Alg. II, 1865, p. 30. 
Diam. cell. 6-7 M; diam. colon. 16 M. 
Hub. Green Lake. 



298 W. AND G. S. WEST 

Colonies small and compact, consisting of eight or sixteen regularly arranged 
cells. Cell- contents of a brilliant blue- green colour and granulose. 

Genus MICROCYSTIS Kiitz. 

82 Microcystis stagnates Lemm. 

in Forschungsber Biol. Stat. PI on. X., p. 150. Polycystis pallida Lemm. P. elongata 
W. & G. S. West. 

Diam. cell. 1.7-1.8 n. 

Hob. Coast Lake. 

RIVULARIACEJ; 

Genus CALOTHRIX Ag. 

83 Calothrix epiphytioa W. & G. S. West 

" Welw. Afric. Freshw. Alg.," Journ. Bot. 1897, p. 240. 
Crass, fil. ad bas. 7.5-8 M ; crass, trich. ad bas. 5.4-5.8 M. 
Hob. Lake at Hut Point. 

84 Calothrix sp. (PI. XXV, Figs. 55-57) 

C. strato tenui submembranaceo serugineo-coaruleo ; filis subbrevibus dense 
intricatis et contortis, paullo attenuatis sed non in pilum productis; trich omatibus 
cceruleis, gradatim attenuatis, cellula terminali obtusa ; cellulis subtorulosis, rotundato- 
rectangularibus, diametro trichomatis paullo brevioribus vel nonnunquam duplo 
brevioribus; heterocystis basilaribus singulis et inflate- hemisphsericis. 

Crass, fil. ad. bas. 10-12 M; crass, trich. ad bas. 8-9.6 M. 

Hob. Blue Lake. 

It is highly probable that this is as yet an unnamed species of Calothrix, but we do 
not feel justified in expressing a definite opinion until more material has been examined. 
It occurred in very small thin patches, of a most intense blue- green colour, among 
the tough sheets of Phormidium glaciale. The filaments were much contorted, densely 
intricate, and quite without any definite arrangement in the stratum. The attenuation 
was slight, and none of the plants showed any signs of hair- like apices. Thesubtorulose 
cells are also a feature of the trichomes. 



PLATE XXIV 



PLATE XXIV 

FIGURES 1-2. Ulothrix cequalis Kiitz. x 500. 

FIGURES 3-4. Ulothrix cequalis Kiitz. forma, x 500. 

FIGURES 5-7. Ulothrix tenerrima Kiitz. forma untarctica. 5 and 6, x 500 ; 7, 

x 1000. py, pyrenoid. 
FIGURES 8-9. Prasiola crispa (Lightf.) Menegh. 8, thallus, natural size.- 9, cells of 

small portion of thallus, x 500. 
FIGURES 10-11. Prasiola antarctica Kiitz. 10, thalli, natural size; 11, cells of 

small portion of thallus, x 500. 
FIGURES 12-14. Prasiola crispa (Lightf.) Menegh. Hormidium- stage, from vicinity 

of Cape Royds. x 500. 

FIGURES 15-18. Prasiola crispa var. aspera var. n. x 500. 
FIGURES 19-24. Chlamydomonas intermedia Chodat forma antarctica. x 500. py, 

pyrenoid ; st, pigment spot. 23 and 24 show stages of division. 

FIGURES 25-29. Chlamydomonas subcaudata Wille. x 500 n, nucleus; py, pyre- 
noid ; st, pigment spot. 28 and 29 show stages of division. 
FIGURES 30-39. Pleurococcus dissectus (Kiitz.) Nag. x 500. 
FIGURES 40^14. Pleurococcus frigidus sp. n. x 500 n, nucleus; py, pyrenoid; 

s, small granules of starch. 
FIGURES 45-46. Pleurococcus pachydermusLagerli. x 500. n, nucleus ; py, pyrenoid. 

46 shows escape of eight daughter- cells (gonidia) from old mother- cell. 
FIGURES 47-48. Pleurococcus pachydermus forma stipitata. x 500. 
FIGURES 49-51. Pleurococcus antarcticus sp. n. x 500. ol, globules of fatty oil. 
FIGURES 52-54. Pleurococcus antarcticus forma robusta. x 500. 



Brit. Antarct. Exped. 1907-9. 



Vol. 1. Plate XXIV. 



W. & G. S. WEST: FRESHWATER ALG/E. 



OO 



-~ 

98 88 GO 
IB 808 00 

CDCD SS QQ \jQ 



u . 




G. S. WEST. ad. nat. del. 



PLATE XXV 



PLATE XXV 

FIGURES 55-57. Calothrix sp. x 500. 

FIGURES 58-67. Nostoc antarctica sp. n. 58, young thalli, nat. size ; 59, fully grown 
thalli, nat. size; 60, margin of adult thallus showing the dense crowding of the 
trichomes, x 500 ; 61-65, portions of isolated trichomes showing seriate hetero- 
cysts, x 500 ; 66, very young thallus, x 500 ; 67, portion of trichome with five 
spores (sp.}, x 500. 

FIGURES 68-69. Lyngbya shackletoni sp. n. x 500. 

FIGURES 70-71. Lyngbya murrayi sp. n. 70, x 500; 71, x 1000. 

FIGURE 72. Lyngbya erebi sp. n. x 1000. 

FIGURE 73. Phormidium glaciate sp. n. x 1000. 

FIGURES 74-75. Phormidium antarcticum sp. n. x 1000. 

FIGURE 76. Phormidium fragile (Menegh.) Gom. forma tennis, x 1000. 

FIGURES 77-85. Phormidium autumnale (Ag.) Gomont. Various forms, x 500. 

FIGURES 86-90. Oscillatoria producta sp. n. 86, x 500 ; 87-90, x 1000. 

FIGURES 91-94. Oscillatoria subproboscidea sp. n. x 500. 

FIGURES 95-97. Oscillatoria deflexa sp. n. 95, twisted bundle of trichomes, x 100 ; 
96 and 97, single trichomes, x 1000. 



Brit. Antarct. Exped. 1907-9. Vol. 1. Plate XXV. 

W. & G. S. WEST: FRESHWATER ALGAE. 




G. S. WEST. ad. nat. del. 



PLATE XXVI 



BRIT. ANTARCT. EXPED. 1907-9. VOL. I. 



PLATK XXVI 

FIGURE 98. Oscillatoria terebriformis Ag. forma tennis, x 500. 
FIGURES 99-101. Oscillatoria priestleyi sp. n. 99, x 500 ; 100 and 101, x 1000. 
FIGURE 102. Oscillatoria amphibia Ag. var. robusta var. n. a, x 500 ; b, x looo. 
FIGURES 103-106. Asterocystis antarctica sp. n. 103-105, x 500; 106, x 1000. 
FIGURES 107-114. Microcystis chroococcoidea sp. n. x 500. 

FIGURES 115-120. Tropidoneis laivissima sp. n. 115-118 and 120, x 500; 119, 
x 1000. Fig. 117 is one of the deformed valves not infrequent in Clear Lake. 
Fig. 115 is the girdle view, and Fig. 120 the end view of a frustule. 
FIGURES 121-124. Navicula muticopsis V. Heurck. x 1500. 
FIGURE 125. Navicula glaberrima sp. n. x 1000. 

FIGURES 126-127. Achnanthes brevipes Ag. var. intermedia (Kiitz.) Clove, x 1500. 
FIGURE 128. Fragilaria tenuicollis Heib. var. antarctica var. n. x 1500. 
FIGURE 129. Navicula murrayi sp. n. x 1500. 
FIGURE 130. Navicula murrayi var. elegans var. n. x 1500. 
FIGURE 131. Navicula muticopsiforme sp. n. x 1500. 
FIGURE 132. Navicula peraustralis sp. n. x 1800. 
FIGURES 133-134. Navicula cymatopleura sp. n. x 1500. 
FIGURE 135. Navicula globiceps Greg, x 1500. 
FIGURES 136-138. Navicula shackletoni sp. n. x 1500. 
FIGURES 139-141. Navicula shackletoni var. pellucida var. n. x 1500. 



Brit. Antarct. Exped. 1907-9. 



Vol. 1. Plate XXVI. 



W. 6" G. S. WEST: FRESHWATER ALG/E. 



107. 108. ilO. 




12.8. 



135. 



140. 



G. S. WEST. ad. nat. del. 



BRITISH ANTARCTIC EXPEDITION 1907-9 

UNDER THE COMMAND OF SIR E. H. SHACKLETON, C.V.O. 



REPORTS ON THE SCIENTIFIC INVESTIGATIONS 




VOL. II 

BIOLOGY 

EDITOR - - JAMES MURRAY 

PART I 

MOLLUSC A 

BY CHAS. HEDLEY 

(AUSTRALIAN MUSEUM, SYDNEY) 



(WITH OXE PLATE) 



LONDON 
PUBLISHED FOR THE EXPEDITION BY WILLIAM HEINEMANN 

21 BEDFORD STREET, W.C. 

1911 

ISSUED SEPARATELY, APRIL, 1'Jll 
PRICE l.v. 6d. NET 



REPORTS ALREADY ISSUED 

VOL. 1. PART I., ON COLLECTING AT CAPE UOYDS 

BY JAMES MURRAY PRICE Is. (id. NET 

VOL. I. PART II., ON MICROSCOPIC LIFE AT CAPE ROYDS 

BY JAMES MURRAY PRICE 5s. Od. NET 

VOL. I. PART 111., ANTARCTIC ROTIFERA 

BY JAMES MURRAY PRICE 5s. Od. NET 

VOL. I. PART IV., MUSCI 

BY JULES CARDOT P 1(KE is. Od. NET 

VOL. I. PART V., TARDIGRADA 

BY JAMES MURRAY PR,CE 10s. Od. NET 

VOL. I PART VI., RHIZOPODES D'EAU DOUCE 

BY EUGENE PEXARD P HIC E 3s. Od. NET 



PART I 

MOLLUSCA 

BY CHARLES HEDLEY 

(Prepared by permission of the Trustees of the Australian Museum, Sydney) 

(With Plate I.) 



EDITORIAL NOTE. Mr. Hedley has asked me to preface his report with field notes on the species. The 
Mollusca were collected in two localities, within a short distance of one another at Cape Royds, yet of 
totally difl'erent character. The "Bay" is a very small shallow inlet, lying to the east of our camp, 
between Gape Royds and Cape Barne. It appears to be little affected by currents, or even by storms, for 
the ice forms early there and stays late. Wherever it has been examined at depths of more than two 
fathoms, the bottom resembles the bed of a lake in being covered with a fine tenacious black silt. In 
consequence of the early formation of the winter ice in the Bay we were able to dredge there as early as 
April, and frequently afterwards till July. Then this dredging-ground was lost, and no more work was 
done in the Bay till February, when the ice had gone out of part of the Bay. The Bay is only free from 
ice for a few weeks in summer, perhaps in cold seasons not at all. The dredging was done at depths of 
between seven and twenty fathoms. 

The " Sound " is a situation of very different character. The part where most of the dredging was done 
is at the mouth of the Bay, where the bottom begins to slope down into deeper water. A strong current 
prevails there, as indicated by Brocklehurst's Current Indicator, and by the effect on our dredging lines. 
The bottom is free from mud, even in shallow water close by the cliffs of Cape Royds. At a depth of less 
than twenty-five fathoms there are many pebbles and boulders of keny te and other rocks. At greater depths 
no pebbles are found, the dredge bringing up only living or dead organisms, which apparently grow so 
densely crowded together that the dredge never touches the bed of the sea, of whatever materials composed. 

The molluscan fauna of these two localities differs greatly. In the Bay the dominant forms are 
Neobuccinum, Anatina, Pecten, and Yoldia. The delicate Trophon is not rare, and there are many small 
and even microscopic molluscs, both Gastropods and Lamellibranchs. 

Neobuccinum eatoni appears to surpass all the other large species in abundance, but this may be due to 
its activity and voracity. It comes readily to any bait which is put down for even a short time. No trap 
or dredge is needed to get them. They hold tight on to the bait and come up with it. They must have 
keen senses, of whatever sort, as the crowds which sometimes covered the bait must have collected from 
some little distance around. The shells of Pecten colbecki are very abundant, but the animal was rarely found 
alive. The first living example was got on June 17, between seven and twelve fathoms. On account of the 
delicacy of its valves it was removed from the bucket and placed on the ice for safety. The temperature 
was low, and it showed its disapproval of its first experience of a temperature ( - 10 F.) forty degrees below 
that to which it was accustomed by snapping angrily for some time. When replaced in the bucket and 
conveyed to the house it seemed none the worse, and again snapped when exposed to the air. It lived for 
many hours, under the influence of a weak narcotic (Eucaine), thus showing a tenacity of life unusual 
among Antarctic marine animals, and unexpected in this delicate Pecten. A very small specimen was 
dredged on a stony bottom, at a depth of only two fathoms, close to the shore. 

The shells of Anatina are very abundant and of large size, but only very small examples were got alive. 
BEIT. ANTARCT. EXPED. 1007-9. VOL II. PART 1, ISSUED APRIL 1911 A 



/3 



2 CHARLES HEDLEY 

It may be that the living molluscs are too firmly embedded in the mud to be detached by the dredge. 
The mud which has been stirred up by the dredging, when allowed to settle in the bucket, is so tenacious 
that it is with difficulty that a half -buried Anatina shell can be pulled out. 

The living Yoldia, as well as shells, are very plentiful. 

In the Sound the large molluscs are entirely different. The conspicuous forms of the muddy bottom 
are absent, except for an occasional empty shell of Anatina or Yoldia, and instead we find Lima, Tritoniella, 
Philobrya, Nation, and others. 

Only the Lima and Philobrya are at all abundant ; the other conspicuous forms are rather rare. The 
Lima is constantly present at depths of twenty-five to eighty fathoms. Very commonly the animal is 
embedded in a sponge, usually in one of the softer horny kinds. 

Some sorts of Opisthobranchs (Tectibranchs) were obtained both in the Bay and the Sound, but the 
best specimens were got in the mouths of the Big-head Fishes (Trematomus). 

A comparison of our collection with that of the Discovery Expedition is instructive for the light it 
throws on local distribution in the Ross Sea area. 

Leaving out of account the Opisthobranchs, which are not yet fully worked out for our Expedition 
(although Mr. Hedley records a few species), our collection includes ten Lamellibranchs and twenty 
Gastropods, against fourteen Lamellibranchs and twenty-five Gastropods obtained by the Discovery. Of 
these, seven Lamellibranchs and twelve Gastropods are common to the two collections. The Discovery had 
thus twenty species (seven Lamellibranchs and thirteen Gastropods) which were not in our collection, and 
we had eleven species (three Lamellibranchs and eight Gastropods) which were not in the Discovery collection. 

This amount of difference between'collections made at stations only twenty miles apart is very considerable. 
It is more important that one of our most abundant shallow-water species ( Yoldia eightsi) did not occur at 
all at Hut Point (the Discovery winter quarters). 

Of the species which occurred in our collection and not in that of the Discovery, one Gastropod and 
one Lamellibranch were got by the Sotithern Cross far to the north of our position, and one Gastropod 
was collected by the Belgka. 

Seven of our species (two Lamellibranchs and five Gastropods) are described by Mr. Hedley as new to 
science. J. M. 



SIR ERNEST SHACKLETON has done me the honour to hand me for report the 
collection of mollusca accumulated by the British Antarctic Expedition of 1908 
under his command. This consists chiefly of specimens dredged by Mr. James Murray 
through holes and cracks in the ice around Cape Royds. In this neighbourhood the 
slope of Mount Erebus continues under the sea, so that shallow and deep stations 
lie near together. 

For several feet below the surface all life is destroyed by the floating ice, but 
below its influence a fauna flourishes which, for the latitude, may be considered 
luxuriant. The bottom in shoaler depths is described by Mr. Murray as carpeted 
with a matted growth of weed and sponge, beyond which was an area of soft 
black rnud. 

Salient characters of the collection, which strike an observer accustomed to 
tropical and temperate faunas, are the great chemical erosion which the shells have 
suffered, and against which some, Yoldia for instance, have developed a dense 
epidermis. A large proportion are frail and thin. 

No great number of novelties could be anticipated from this collection, for the 
Expedition's field of examination had already been exploited by the Discovery; 



MOLLUSC A 3 

while the area searched by the Southern Cross was sufficiently near to infringe 
upon their prospects. But viewing this collection as a control upon its predecessors 
and as an intense r study of the fauna of McMurdo Sound, we gain from it a better 
idea of the distribution of the Antarctic marine fauna. 

L AMELLIBRA NCHIATA 

Yoldia eif/htsi, Couthouy 

Smith, Report Collect. Southern Cross, 1902, p. 211. 
Alive, abundant, from five to thirty fathoms. 

Adacnarca nitens, Pelseneer 

Smith, Report Nat. Antarctic Expecl, Lamell., 1907, p. 5, Plate III., Figs. 6-6c. 
Alive, twelve specimens from eighteen to twenty fathoms. 

Philobrya limoides, Smith 

Smith, Report Nat. Antarctic Exped., Lamell., 1907, p. 4, Plate III., Figs. 2-26. 
Alive, abundant from ten to eighty fathoms. 

Pecten colbecki, Smith 

Smith, Report Nat. Antarctic Exped., Lamell., 1907, p. G, Plate III., Figs. 9-9a. 
Several living specimens, largest about 83 x 79 mm., from two to twelve fathoms. 
Several double and many single and broken valves from ten to twelve fathoms. 

Lima hodgsoni, Smith 

Smith, Report Nat. Antarctic Exped., Lamell., 1907, p. G, Plate III., Figs. 8-86. 
Alive, abundant from ten to eighty fathoms. 



Anatina elliptica, King and Broderip 

Smith, Report Nat. Antarctic Exped., Lamell., 1907, p. 1. 
Alive, abundant from seven to thirty fathoms. 

Thracia meridionalis, Smith 

Smith, Report Nat. Antarctic Exped., Lamell., 1907, p. 1. 

Alive, a few specimens, mostly broken, from ten to eighty fathoms. 

Gardita astartoides, Martens 

Smith, Report Nat. Antarctic Exped., Lamell., 1907, p. 2. 
A couple of broken valves from ten to twenty fathoms. 



4 CHARLES HEDLEY 

Kellia nimrodiana, sp. nov. (Plate L, Figs. 1-4) 

Shell, subi-homboidal, slightly inequilateral, thin, inflated, glossy, white. 
Sculpture : fine irregular concentric growth lines without radials. Hinge line 
rather straight, umbo tumid, prodissoconch conspicuous. Ligament internal, right 
valve with a prominent cardinal knob, left valve with a feeble ridge, no laterals. 
Length, 3 '8 ; height, 4'25 ; depth of conjoined valves, 3'1 mm. 

Fifteen specimens alive from ten to twenty fathoms off Cape Royds. 

Solecardia (Antarctica, sp. nov. (Plate L, Fig. 5) 

Shell small oblong compressed, semi-transparent, gaping at each end, polished, 
with a few radial stria;. Length, 1'25 ; height, 0'8 mm. 

Five specimens alive from ten to twenty fathoms off Cape Royds. 

GASTROPODA 

Lepeta antarctica, Smith 

Smith, Hep. Nat. Antarctic Exped., Gast., 1907, p. 12, Plate II., Figs. 11-1 la. 
Two specimens alive from eighteen to eighty fathoms. 

Valvatella refulyens, Smith 

Smith, Rep. Nat. Antarctic Exped., Gast., 1907, p. 11, Plate II., Fig. 7. 
Abundant alive from eighteen to eighty fathoms. 



crebrilirulata, Smith 

Smith, Rep. Nat. Antarctic Exped., Gast., 1907, p. 11, Plate II., Fig. 9. 
Abundant alive from ten to twenty fathoms. 

Lacuna macmurdensis, sp. nov. (Plate L, Fig. G) 

Shell small, turbinate, thin, and loosely coiled. Colour, pale buff. Sculpture : 
three strong peripheral keels, beneath which are two smaller threads. Base 
flattened, angled at the margin of a rather wide spiral umbilicus. Major 
diam., 3'5 mm. 

A single broken specimen from ten to twenty fathoms oft' Cape Royds. 

This appears to represent L. noto-rcadcitsis, Melvill and Standen,* from the opposite 
coast of Antarctica. It differs by having the shoulder of the whorl sloping instead 
of flat, fewer and more prominent keels, and a broad smooth area between the lowest 
keel and the umbilical margin. 

* Melvill and Standen, Trans. Roy. Soc. Edinburgh, xlvi., 1907, p. 181, Plate, Figs. 3-3. 



MOLLUSCA 5 

Rissoa adarensis, Smith 

Smith, Hep. Nat. Antarctic Exped., Gast., 1907, p. 8, Plate II., Fig. 2. 
Six shells from ten to twenty fathoms. 

Rissoa fraudulenta, Smith 

Smith, Rep. Nat. Antarctic Exped., Gast., 19Wrp- 9, Plate II., Fig. 3. 
One shell from ten to twenty fathoms. 

Rissoa gelida, Smith 

Smith, Rep. Nat. Antarctic Exped., Gast., 1907, p. 9, Plate II., Fig. 5. 
Alive, twenty specimens from ten to twenty fathoms. 

Rissoa glacialis, Smith 

Smith, Hep. Nat. Antarctic Exped., Gast., 1907, p. 9, Plate II, Fig. 4. 
Two shells from ten to twenty fathoms. 

Capidus subco'inpressus, Pelseneer 

Pelseneer, Result voy. Belgica, Moll., 1903, p. 20, Plate V., Figs. 52-54. 
One alive from " Bay, 2/09." This is an addition to the fauna of this 
quadrant of Antarctica. 

Lovenella antarctica, Smith 

Smith, Rep. Nat. Antarctic Exped., Gast., 1907, p. 10, Plate II., Figs. G-G. 
A single shell. (Loc. Sound, twenty-five to fifty fathoms, " 6/7/08 ") 

Lovenella austrina, sp. nov. (Plate I., Fig. 7) 

Shell small elongate-conical, thin and semi-transparent. Colour pale buff. 
Whorls ten, including a smooth helicoid protoconch of a whorl and a half. 
Sculpture : three slender spiral keels on each whorl, of which one margins the lower 
suture, one is peripheral, and one intermediate ; on the last whorl an incipient 
thread appears above the periphery and another on the base. These are beaded 
by the passage of fine numerous perpendicular irregular riblets which also cancellate 
the interspaces. Aperture subcircular, angled above, notched below by a very 
short perpendicular canal, columella thickened, arched, outer lip thin. Length, 8 ; 
breadth, 2 -5 mm. 

Twenty specimens alive from ten to twenty fathoms off Cape Royds, June 1908. 

In size and shape closely resembling L. antarctica, but differing by fewer more 
delicate spirals and thinner shell. Cerithium charcoti, Lamy,* from Booth Wandel 
* Lamy, Exped. Charcot, Moll., 1900, p. ], Plate I., Fig. 1. 



6 CHARLES HEDLEY 

Island, seems to be related, but is broader and has an extra keel on the shoulder of 
the whorl. 

Thesbia, innocent, Smith 

Smith, Rep. Nat, Antarctic Exped., Gast., 1907, p. 4, Plate I., Figs. 1-lfc. 
A single shell, Sound, twenty-five to fifty fathoms. 

Vermici/htria murmyi, sp. nov. (Plate I., Fig. 8) 

Shell solitary, small, solid, compactly coiled in about three discoidal whorls, the 
last rising free for almost a third of a revolution. Sculpture : on the upper surface 
three prominent equidistant spiral ridges parted by deep interstices. Aperture 
circular, slightly lobed. Diameter of coil, 4'25 ; diameter of aperture, 1'65 mm. 

Several specimens, mostly young and fragmentary, from sixty to eighty fathoms 
off Cape Royds. 

This species is closely related to V. icaitei, Hedley,* from off Sydney, but is 
larger, more solid, has the spirals closer, and does not seem to develop variceal rings. 
It is named in honour of Mr. James Murray, the Biologist of the Expedition. 

Odostorniopsis major, sp. nov. (Plate I., Figs. 9-10) 

Shell ovate-conical, subperforate, very thin. Whorls four and a half, rounded, 
parted by deep sutures. Colour, white. Sculpture : on the second whorl half a dozen 
spiral punctate lines, the remainder smooth, save for microscopic incremental 
threads. Aperture roundly pyriform, outer lip thin, effuse anteriorly. Columella 
flattened, expanded over the axial perforation and then sharply bent. Length, G ; 
breadth, 4 mm. 

Type labelled " Bay, July, 1908 " ; another specimen eighteen to twenty fathoms, 
and two more from sixty to eighty fathoms off Cape Royds. All alive. 

The genus Odostomiopsis was proposed by Thiele t for the reception of two species 
from Kerguelen. To this genus should probably be referred Admete (?) limmmformis , 
Smith,;}: also from Kerguelen. The appearance of the species suggests to me that 
possibly it may be, like Stilifer, a parasite upon Echinoderms. 

Neobuccinum eatoni, Smith (Plate I., Figs. 11-12) 

Smith, Rep. Nat. Antarctic Exped., Gast., 1907, p. 1. 

Alive, abundant from five to sixty fathoms. The expeditions led by Dr. Bruce 
and Dr. Charcot respectively have recently extended the range of this species to 

* Hedley, Mem. Austr. Mus., IV., 1903, p. 346, Fig. 72. 

f Thiele, in Martens Gast. d. deut. Tiefsee Exped., 1898-1899, p. 68 (1903); not Odontomopsis, 
Blanckenhorn, Beitr. z. Geol. Syriens, 1890. 

t Smith, Phil. Trans. Roy. Soc. Lond., clxviii., 1879, p. 172, Plate IX., Fig. 4. 
Wilton, Zoological Log, Scotia, 1908, p. 2, Plate XXIV., Fig. 74. 



MOLLUSCA 7 

the opposite coast of the continent. The radula has lately been redescribed by 
Professor Thiele.* 

All the adult specimens have the apex eroded, but the youngest examples show 
it (Fig. 11) to be dome-shaped and paucispiral. 

The collection contains a quantity of molluscan ova labelled "7-20 fath., 
June 1908," occurring singly or plastered together in masses. From its size and 
abundance these can with some confidence be asertbed to N. eatoni. Each capsule 
(Fig. 12) is a hemispherical orange boss about a third of an inch in diameter and 
surrounded by a marginal membrane. The ova of Cominella densisculpta, Martens, f 
seem to be similar. 

Amciuropsis rossiana, Smith 

Smith, Rep. Nat. Antarctic Exped. ; Gast., 1907, p. 5, Plate I., Figs. 6-Ga. 
Three dead and broken shells from five to twenty fathoms. 

Naticct yrisea, von Martens 

Watson, Rep. ChalL, Zool, XV., 1886, p. 432., Plate XXVIII, Fig. 5 ; Thiele, 
Deut. Tiefsee Exped., VII., p. 64, Plate VIIL, Fig. 44. 
Two dead shells from ten to twenty fathoms. 

Lamellaria mollis, Smith 

Smith, Report Coll. Southern Cross, 1902, p. 205, Plate XXIV., Figs. 19-21. 
Two small specimens from ten to eighty fathoms. 

Trophon shaMetoni, sp. nov. (Plate I., Fig. 13) 

Shell biconical, large, and thin. Colour dead white. Whorls five, and a subulate 
smooth protoconch, turreted, rather rapidly increasing. Sculpture : broad imbricate 
thin lamellae, spreading from the base upwards, produced above in hollow curled 
spouts which crown the whorl, thence continuing to the suture in a low oblique 
limb. On the last whorl there are from twelve to fourteen lamellae, which continue 
in a diminishing series up the spire. Interlamellar spaces are scored by fine spiral 
striae and more conspicuous scratches in the line of growth. Aperture ample, ovate. 
Canal rather short, screwed to the left, externally with imbricating hoods formed by 
bases of successive lamellae. Specimen figured, 26 mm. long, 18 mm. broad. A 
larger example is 33 mm. long and 23 mm. broad. 

Ten specimens, several alive, from seven to twenty fathoms off Cape Royds. 

This appears to be closely related to T. coulmanensis, Smith, \ from which it 

* Thiele, Deut. Tiefsee Exped., VII., p. 1C8, Plate IX., Fig. 57. 

t Martens and Pfeff'er, Jahrb. Hamb. wiss. Anstal. III., 188C, p. 73, Plate I., Fig. P.. 

+ Smith, Discovery, Moll., p. 2, Plate I., Figs. 4-46. 



8 CHARLES HEDLEY 

differs by greater size, broader shape, more compact spire, shorter and more bent 
canal. Seeing the variability of the genus, I distinguish this as new with some 
diffidence, but the series before me is quite uniform, and T. coulmanengig is reported 
from much deeper water, viz., 100 fathoms. As the handsomest novelty in the 
collection, it is dedicated to the intrepid leader of the Expedition. 

The apex is of the pattern figured for T. coulmanensis. In aged specimens several 
of the spire whorls are usually destroyed by erosion. The interlamellar spaces of the 
last whorl are usually packed with sponges and diatoms, thus providing the shell 
with an additional overcoat, a possible defence against chemical erosion. 

Trophon lonystaffi, Smith 

Smith, liep. Nat. Antarctic Exped., Gast., 1907, p. 3, Plate I, Figs. 3-3cZ (Plate I., 
Fig. 14). 

Three specimens alive from twenty to eighty fathoms. 

As Mr. Smith observed, the upper whorls are more closely ribbed than the last. 
This causes much dissimilarity between younger and older shells. An immature 
shell 25 mm. in length is here figured. 

Philine apertissima, Smith 

Smith, Rep. Coll. Southern Cross, 1902, p. 208, Plate XXIV., Fig. 23. 
Twenty living specimens from ten to twenty fathoms. 

Philine antarctica, Smith 

Smith, Hep. Coll. Southern Cross, 1902, p. 208, Plate XXIV., Fig. 1. 
Six living specimens from ten to twenty fathoms. Others from stomach of fish 
(Big-head, Trematomus). 

Clione antarctica, Smith 

Smith, Rep. Coll. Southern Cross, 1902, p. 210, Plate XXV., Figs. 7-8. 
Two specimens from surface near Cape Royds. 

Limacina antarctica, Woodward 
Smith, Rep. Coll. Southern Cross, 1902, p. 209. 

A few specimens from surface near Cape Royds. Their shells were dissolved by 
the formed in which they were placed. 

Tritoniella belli, Eliot 

Eliot, Rep. Nat. Antarctic Exped., Nudibr., 1907, p. G, Figs. A, B. 
Several specimens from twenty to fifty fathoms off Cape Royds. Some coils of 
molluscan ova marked " 15-25 fath., 7/08 " probably belong to this species. 



PLATE 



BtUT. ANTAKOT. KXPEU. l!H)7-!>. VOL. II. 



PLATE I 



PAOE 



FIGURES 1-4. KelUa nimrodiana, Hedley . . 4 

FIGURE 5. Solecardia antarctica, Hedley . 4 
FIGURE 6. Lacuna macmurdensis, Hedley . 

FIGURE 7. Lovenella austrina, Hedley . . 5 

FIGURE 8. Vermicularia murrayi, Hedley fi 

FIGURES 9-10. Odostomiopsis major, Hedley . 6 

FIGURE 11. Apex of Neobuccinum eatoni, Smith 6 

FIGURE 12. Eggs of Neobuccinum eatoni, Smith 7 

FIGURE 13. Trophon shackletoni, Hedley . . 7 

FIGURE 14. Trophon longstaffi, Smith. 8 



BRIT. ANTARCT. EXPED. 1907-9 

HEDLEY: MOLLUSCA 



VOL. II. PLATE I 






10 






9 




11 




14 





Miss West del. 



, BRITISH ANTARCTIC EXPEDITION 1907-9 

UNDER THE COMMAND OF SIR E. H. SHACKLETON, C.V.O. 



REPORTS ON THE SCIENTIFIC INVESTIGATIONS 



VOL. II 

BIOLOGY 

EDITOR - - JAMES MURRAY 
PART II 

ANTARCTIC FISHES 

BY EDGAR R. WAITE, F.L.S. 



(WITH OXE PLATE) 



LONDON 
PUBLISHED FOR THE EXPEDITION BY WILLIAM HEINEMANN 

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1911 

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PART II 

PISCES 

ANTARCTIC FISHES 

BY EDGAR R. WAITE, F.L.S. 

Curator, Canterbury Museum, New Zealand 
(With Plabe II.) 

THE collection of fishes placed in my hands for report by Sir Ernest Shackleton is 
quite a small one, comprising five species only. 

Of these, four were described by Dr. Boulenger from the proceeds of the 
Southern Cross Expedition.* When examining this latter collection, Boulenger 
separated off, on an osteological character, certain Nototheniids under the name 
Trematomus : four species were referred to this genus, namely, T. newnesi, 
T. borchgrevinki, T. hansoni, and T. bernacchii, all from the Antarctic, and new, whilst 
fifteen were retained as Notothenia, two only of which were taken in Antarctica, 
namely, N. nicholai and N. coriiceps. 

Reporting on the fishes of the National Antarctic Expedition, 1901-1904, taken 
on the voyage of the Discovery, Boulenger t records three of the four known 
species of Trematomus, that is, T. borchgrevinki, T. hansoni, and T. bernacchii, and 
four species of Notothenia, two of which were new, namely, N. scotti and N. hodgsoni. 

Among the collection obtained by the Expedition Antarctique Frangaise, 1903- 
1905, Professor VaillantJ found eight species of Notothenia, but not a single 
species of Trematomus. 

In the absence of the reports by Dollo on the fishes of the Belgica Expedition, 
and by Lonnberg on those of the Swedish South Polar Expedition, I cannot make 
further comparison, and the want of the latter paper makes it impossible for me to 
specifically determine, at present, the single example of the genus Artedidraco taken 
on the Expedition. 

Lonnberg has described two species of this genus, namely, Artedidraco mirus and 

* Boulenger, SoutJi&rn Cross Exped. Report, 1902, Pisces, p. 174. 
t Boulenger, Zoology, Nat. Antarct. Exped., Fishes, 1907. 
J Vaillant, Exped. Antarct. Frang., 1906, Poissons. 

BEIT. ANTARCT. EXPED. 1907-9. VOL. II. PART 2, ISSUED APRIL 1911 C 

5 



12 EDGAR H. WAITE 

A. skottsbergi, but their distinguishing characters are not at present known 
to me. 

The four species of fishes which have been identified are : 

Trematomus bernacchii, Boulenger. 
Trematomus hansoni, Boulenger. 
Trematomus newnesi, Boulenger. 
Pleuragramma antarcticum, Boulenger. 

The gill cavities of the two first named yielded small crustaceans, while parasitic 
forms were found upon T. bernacchii. All were submitted to Dr. Charles Chilton, 
and he has kindly identified them as follows : 

Euphausiacea. 

A single damaged specimen, not identifiable. 
Isopoda. 

Antarcturus franklinii, Hodgson, one imperfect specimen. 
Amphipoda. 

Orcliomenopsis rossii, Walker, several specimens. 

Orchomonella pinguides, Walker, several specimens. 

Harpinia obtusifrons, Stebbing, one specimen. 

Leptamphosus novce-zealandice, G. M. Thomson ( = Oradarea longimana, 

Walker), one specimen. 
Copepoda. 

Calanus, sp., one specimen. 

Parasitic Copepod, several specimens from T. bernacchii. 

TREMATOMUS, Boulenger, 1902 

Trematomus bernacchii, Boulenger 
Keport Southern Cross Collections, 1902, p. 181, Plate XIV. 

This species is characterised by the narrow interorbital space, which is also devoid 
of scales, with the exception of a few disposed in a single median longitudinal row. 

In most of the examples preserved the upper half of the first dorsal fin is black or 
blackish, as described, though in some there is no trace of such ; the other fins bear 
no markings. In addition to the coloration described by Boulenger, a few specimens 
possess a black blotch on the upper part of the opercle. The ground colour in 
preservative is reddish brown, but the red tint is probably more pronounced in life, 
for Mr. Murray has recorded the colour of some examples as " strongly ruddy," 
" light ruddy," &c. 

Specimens examined were in full ripe roe when taken on August 25, 1908 ; the 
ova are bright yellow in colour and very large, measuring 3 '8 mm. in diameter. 



PISCES: ANTARCTIC FISHES 13 

Eighteen specimens were preserved, of which the largest measures 239 mm. in 
length ; they were localised as follows : 

Nos. 1-3. The Bay, Cape Eoyds, taken in trap 18 fathoms, May 20, 1908. 

No. 4. The Bay, Cape Royds, collected by Mawson, May 11, 1908. 

Nos. 5-7. Cape Royds, 30-80 fathoms, August 1908. 

Nos. 8-13. Bay, 10-20 fathoms, June 1908. 

No. 14. The Bay, Cape Royds, dredge '10-20- fathoms, May 12, 1908. 

Nos. 15-18. Sound, 10 fathoms, July 7, 1908. 

I found small examples of Trematomus newnesi in the digestive tracts of some 
of the specimens, and the gills of most of the fishes submitted harboured many 
crustaceans ; these have been submitted to Dr. Charles Chilton, who furnished the 
accompanying report. A Copepod (Calanus, sp.) was found to be parasitic on 
this fish. 

Trematomus hansoni, Boulenger 

Report Southern Cross Collections, 1902, p. 180, Plate XIII. 

The narrow scaly interorbital space and the strongly barred fins are characteristic 
of this species, the markings in the specimens preserved being much more pronounced 
and better defined than portrayed by Boulenger : this feature was recognised as 
distinctive by Mr. Murray, who wrote : " Big-heads with black fins and markings." 
One example has the edge of the preopercle white, continued as a band across the 
occiput. 

Though the ground colour in preservative is brownish, Mr. Murray renders the 
tint in fresh specimens as " dark greenish," " light grey," &c. 

Of six examples submitted the largest measures 249 mm. in length. 

The following are the localities of the specimens : 

Nos. 1-2. In fish-trap, Cape Royds, 25 fathoms, September 1, 1908. 
No. 3 (cut in two). Found on ice beside seal-hole in the Bay, Cape Royds, 
May 1908. 

Nos. 4-6. In fish-trap, Cape Royds, 25 fathoms, August 25, 1908. 

Dr. Charles Chilton has examined the crustaceans found in the gill cavities oi 
these fishes, and his report is printed herewith. 

Trematomus newnesi, Boulenger 
Report Southern Cross Collections, 1902, p. 177, Plate XI. 

With two exceptions all the examples referred to this species are small. The 
largest measures 137 mm. in length ; most of the others are much smaller. 

The relatively wide space between the eyes is a character common also to 
T. borchyrevinki, Boul., but T. neivnesi is distinguishable from that species by the 
strongly marked and tubular lateral lines. 

The specimens placed in my hands agree generally with the description of this 



14 EDGAR R. WAITE 

species, but the tail is truncate and not rounded, the principal outer rays being as 
long as the inner ones. The pores on the upper surface of the head are, on the 
whole, as illustrated, but the median pore is placed more forward than shown, so that 
the five interorbital pores form a cross, the median pore being nearly in the axis. 

Young examples have much the appearance of Notothenia hodysoni, Boul.,* 
and were so tentatively named by Mr. Murray. I have, however, examined the 
scapular arch, and find them to possess the distinguishing character of Trematomus ; 
moreover, the vertebrae number fifty-four, which is the number given by Boulenger 
for T. newnesi. 

The localities of the eleven specimens submitted are as follows : 

Nos. 1-6. Backdoor Bay (Priestley), 5-10 fathoms, February 6, 1909. 
No. 7. The Bay, Cape Royds, 5-10 fathoms, May 8, 1908. 

No. 8. " In mouth of Big-head," taken in fish-trap, Cape Royds, 25 fathoms, 
September 1908. 

Nos. 9-11. The Sound, Cape Royds, 10-25 fathoms, July 1908. 

PLEUKAGBAMMA, Boulenger, 1902 

Pleuragramma antarcticum, Boulenger 
Report Southern Cross Collections, 1902, p. 187, Plate XVIII. 

The larger of the two specimens preserved was contained in a bottle with 
examples of Trematomus newnesi, labelled " Backdoor Bay, Priestley, five to ten 
fathoms, February 6, 1909"; the smaller [one was with specimens of T. hansoni, 
August 25, 1908. 

The appearance of the fishes suggests that they were taken from the stomachs of 
a larger fish, or of a seal, all the fins having been more or less digested away. 

The type specimens were also in poor condition, but were sufficiently complete to 
enable an accurate description and figure to be made, as confirmed by Vaillant,t 
who had two excellent examples from the " Francaise " Expedition. He was enabled 
to give a greater range in the number of fin rays, and shows that a rudimentary 
lateral line exists on the hinder part of the body, whereas Boulenger did not find 
trace of such. Vaillant gives detailed descriptions and figures of the scales. The 
scales, though thin, are very adherent, and many of them still remain on our fishes, 
though, as above stated, the digestive process has proceeded to a considerable extent, 
and in the case of the larger example has removed the whole of the hinder part of 
the body. 

One of our fishes appears to have been larger than any previously obtained, 
though its length, except by analogy, cannot be measured ; in its mutilated state it 
measures ] 78 mm. The head, exclusive of the projecting lower jaw, is 55 mm. in length, 

* Boulenger, Nat. Ant. Exped., II., 1907, Fishes, p. 2, Plate II. (I. in text), Fig. 2. 
t Vaillant, Exped. Ant. Fra^aise, 1906, Poissons, p. 48, Figs. 3 and 4. 



PISCES: ANTARCTIC FISHES 15 

and this, contained three and a half times in the length of the body (the proportion 
approximately rendered by Boulenger and Vaillant), would yield a length 
of 193 mm. 

ARTEDIDRACO, Lonnberg, 1908 

Since writing the foregoing I have received the account of the fishes of the 
Swedish South Polar Expedition by Lonnberg.* 

The genus Artedidraco is therein diagnosed, and is represented by two species 
A. mirus and A. skottsbergi. 

The specimen previously referred to as a member of this genus, but which, owing 
to the want of Lonnberg's paper, could not be specifically determined, proves to 
be a species distinct from either of the two previously described forms. It will be 
known as 

Artedidraco shackletoni, sp. nov. (Plate II.) 
B. V. ; D. V. 27 ; A. 20 ; V. I. 5 ; P. 15 ; C. 12 ; L. lat. 19 + 19. 

Head broad, depressed ; body rounded anteriorly, compressed behind. Length 
of head, 2'8 ; height of body, 4'2 ; length of caudal, 37 in the total ; diameter of eye, 
37 ; interorbital space, 10'2 ; and length of snout, 4'1 in the head. 

Mouth large and horizontal ; the maxilla extends to below the middle of the eye, 
and the upper jaw is the longer ; mental barb simple, its length equal to the diameter 
of the eye. The eye is large and prominent and cuts the upper profile ; the nostril is 
in a long tube, placed a little nearer to the eye than to the margin of the preorbital. 
There are a number of large pores with raised rims on the head placed as shown in 
the figure. Opercular spine as in A. mirus. 

The teeth are confined to the jaws and form a narrow band in each ; the edges of 
the lips and the frenum behind the teeth are studded with minute papillae. 

The two anterior spines of the dorsal are abruptly bent near their tips, possibly 
an individual malformation; the second is the longest, its length being 2 '5 in that 
of the head ; the last spine does not quite reach to the second fin. The latter is high ; 
the longest rays are the seventh to tenth, which are subequal, and a little more 
than one-half the length of the head ; the last few rays are rapidly shortened and 
extend to the base of the caudal rays. The anal rays are much shorter than those of 
the dorsal and have their tips free. Anal papilla prominent. The ventrals are widely 
separated and are set quite horizontally ; the fourth ray, which is the longest, is less 
than half the length of the head. The pectoral has a broad base and is rounded ; it 
does not quite reach the origin of the anal. The caudal is feebly rounded, its peduncle 
is much compressed, and its least depth is less than the diameter of the eye. 

The whole fish is scaleless, but is covered with mucus ; the lateral line is formed 
of nineteen pores, which are a continuation of the post-orbital row of pores, and form 

* Lonnberg, Wiss, Ergeb. Schwed. Sudpolar-Exped., V., Zool. I., 1908. 



16 



EDGAR R. WAITE 



a low ridge which extends to below the base of the eighth-ninth dorsal rays ; the 
pores reappear faintly in the mid line above the fifth anal ray, are nineteen in 
number, and do not quite reach the base of the caudal rays. 

Colours. The single specimen preserved is almost colourless, but bears traces of 
darker markings across the back, and spots on the outer caudal rays. 

Length of specimen, 146 mm. It was taken in company with examples of 
Trematomus bernacchii in August 1908, off Cape Royds, at a depth of thirty to eighty 
fathoms. 

The most apparent difference between A. shackletoni and the other members of 
the genus is in the larger number of spines and rays in the dorsal and anal fins, the 
dorsal spines in the other species being but three in number ; some systematists 
would probably regard the distinction as of generic import. 

In other respects the three species are very similar, A. shackletoni being in some 
characters intermediate. By comparison with Lonnberg's figures,* it is found to 
resemble A. skottsbergi in general form, but otherwise more nearly approaches 
A. minis, agreeing in the protuberant eye, the high second dorsal fin, and possibly 
also the coloration. 

The comparative proportions of the three species are shown below : 





A. MIRUS 


A. SKOTTSBERGI 


A. SHACKLETONI 


Dorsal spines and rays 


III. 23-24 


III. 25 


V. 27 


Anal rays 


17 


19 


20 


Head in length 


2-6 


2-8 


2-8 


Height in length 


3-8 


4-7 


4-2 


Eye in head 


3-9 


3-2 


3-8 


Interorbital space in head 


8-7 


10-0 


10-1 


Snout in head 


3-7 


3-6 


4-1 


Caudal in length 


4-2 


3-3 


3-7 



Lonnberg, loc. cit., Plate IV,, Figs. 14 [A. mirus) and 15 (A. skottsbergi). 



PLATE II 



PLATE II 

Artedidraco shackletoni, Waite, page 15. 
Natural size. 



BRIT. ANTARCT. EXPE1). 1907-9 



VOL. II. PLATE II 



WA1TE : PISCES 




c/5 

c 

I 



E. R. Waite del. 



BRITISH ANTARCTIC EXPEDITION 1907-9 

UNDER THE COMMAND OF SIR E. H. SHACKLETON, C.V.O. 

REPORTS ON THE SCIENTIFIC INVESTIGATIONS 



VOL. II 

BIOLOGY 

EDITOR JAMES MURRAY 

PART III 

MALLOPHAGES 

BY PROF. L. G. NEUMANN 

(TOULOUSE) 



(WITH OVE PLATE) 



LONDON 

PUBLISHED FOR THE EXPEDITION BY WILLIAM HEINEMANN 

21 BEDFORD STREET, W.C. 

1911 

ISSUED SEPARATELY, APRIL, 1911 
PRICE Is. 6d. NET 



REPORTS ALREADY ISSUED 

VOL. I. PART I., ON COLLECTING AT CAPE ROYDS 

BY JAMES MURRAY PRICE Is. 6d. NET 

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BY JAMES MURRAY PRICE 5s. Od. NET 

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BY JAMES MURRAY PRICE 5s. Od. NET 

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PART III 

MALLOPHAGA 

PAR PROF. L. G. NEUMANN, TOULOUSE 

(Avec la Planche III) 



EDITORIAL NOTE. The collection of Bird-lice is a very small one, and with a single exception was not 
obtained in the Antarctic region, but on the Albatrosses of the Southern Ocean. 

The one exception, a variety of Philopterus lari, is interesting as the first species found on Maccormick's 
Skua, and also for its occurrence for the first time in the Antarctic. 

As these parasites appear to be extremely rare in the Antarctic it is worth noting that one was 
observed alive on an Emperor Penguin, though unfortunately the specimen was lost among the down. 
The rarity appears to be real, as we carefully examined many penguins of botli species, Skua Gulls, and 
Giant and Antarctic Petrels, and never saw any Bird-lice except these three, two on the Skua and one on 
the Emperor. J. M. 

MALLOPHAGES 

La collection comprend 54 exemplaires, prove nant de 6 oiseaux, dont 3 Diomedea 
exulans, 2 Thalassogeron culminatus, et 1 Megalestris maccormicki, ce dernier seul 
etant reellement antarctique. 

Tons ces Mallophages representent les especes suivantes : 

1. Philopterus lari magnus (Piaget), sur Megalestris maccormicki. 
2. Lipeurus concinnus Kellogg et Chapman, sur 1 Thalassigeron culminatus. 
3. Lipeurus ferox Giebel, sur 1 Diomedea exulans. 
4. Lipeurus hyalinus n. sp., sur 2 Diomedea exulans. 

5. Taschenbergius brews (Dufour), sur les 3 Diomedea exulans et 1 Thalassigercn 
culminatus. * 



1. Philopterus lari (0. Fabricius) (PI. III., figs. 2 a 4) 

1780. Pediculus lari, 0. Fabricius, Fauna groenlandica, p. 219. 

1842. Docophorus lari, H. Denny, Monographia Anophirorum Britannice, p. 89, PI. V., 

fig. 9. 
1844. Philopterus lari, P. Gervais, Histoire naturelle des Insectes : Apteres, iii., p. 337. 

BRIT. ANTARCT. EXPED. 1907-9. VOL. II. PART 3, ISSUED SEPARATELY APRIL 1911 

D 



20 PROF. L. G. NEUMANN 

1871. Docophorus gonothorax, C. G. Giebel, Zeitschrift f. ges. Naturwss., xxxvii. 

p. 450. 

1874. Docophorus congener, C. G. Giebel, Insecta Epizoa, p. 111. 
1880. Docophorus lari v. mayna, E. Piaget, Les Pediculines, p. 112. 

2 $, sur Megalestris maccormicki Saunders, au Cap Royds, par 77 32' lat. S. et 
166 12' long. E., le 1" Janvier 1909. 

Philopterus lari est une espece commune sur nombre d'especes de Larus. Elle 
est indiquee aussi sur Sula alba, Rissa tridactyla policaris, et Creagrus furcatus. 

Les specimens recueillis sur Megalestris maccormicki, les premiers Mallophages 
fournis par cet Oiseau, correspondent a la sous-espece de Piaget, P. I. magnus : 
dimensions plus fortes que dans le type (1 mm. 7 a 1 mm. 8 pour le $), clypeus 
un peu plus allonge et echancre. Les specimens de Piaget provenaient d'un 
Larus atricilla. 

2. Lipeurus concinnus Kellogg et Chapman 

1899. Lipeurus concinnus, V. L. Kellogg et B. L. Chapman, Occasional Papers of the 
California Academy of Sciences, vi., p. 97, PI. VII., fig. 2. 

1 $, sur Thalassogeron culminatus (Gould), au S. de 1'Ocean Indien, par 43 lat. S. 
et 91 30' long. E. 

L'espece a ete decrite par Kellogg et Chapman d'apres des individus pris sur 
Diomedea albatrus en Californie. 

3. Lipeurus ferox Giebel 

1834. Philopterus diomedece, L. Dufour, Ann. de la Soc. entomol. de France, iv., p. 669, 
PI. XXL, figs. 1 et 2. (NonPediculus diomedece Fabricius, Entomologia systematica, 
iv., 1794, p. 421). 

1834. ? Philopterus pederiformis L. Dufour, Ann. de la Soc. entomol. de France, iv., 
p. 676, PL xxi., fig. 4. 

1864. Lipeurus diomedece H. Giglioli, Quart. Journal of Microsc. Science (N.S.), iv., 
p. 19, PL L, figs. 1 et 2. 

1867. Lipeurus ferox C. G. Giebel, Zeitschrift f. ges. Naturwiss., xxix., p. 195. 

1 $, sur Diomedea exulans L., dans 1'Ocean Indien S., au S. de la Tasmanie (avec 
Lipeurus hyalinus Nn., et Taschenbergius brevis (Dufour). 

Kellogg donne pour notes a cette espece : Diomedea melanophrys, D. brachyura, 
D. nigripes (Ocean Pacifique N.), D. albatrus (Californie), et D. exulans. 

L. ferox a ete etabli par Giebel d'apres un individu $ provenant de D. mela- 
nophrys. En en reprenant 1'etude, Taschenberg a decrit comme $ une forme 
que Kellogg regarde comme etant probablement le $ de L. densus Kellogg. La 
? reelle de L. ferox a ete decrite et figuree par Kellogg (1896). La ? de la collection 
antarctique est conforme a la description et a la figure de Kellogg. 



MALLOPHAGA 21 

4. Lipeurus hyalinus, n. sp. (PI. III., figs, la a Id) 

Cette espece est representee par deux lots, recueillis sur Diomedea exulans L., 
savoir : 

5 $ et 3 ?, dans TOcean Indien S., par 40 lat. S. et 47 long. E. ; 4 $, dans TOcean 
Indien S., au S. de la Tasmanie. 

Description. Male. Longueur 4 mm. 05, largeur 1 mm. Corps large, avec des 
bandes incolores, transparentes et quelques petites taches noires. 

Tete subtriangulaire, longue de mm. 9, plus large (0 mm. 87) en arriere des yeux, 
parabolique et incolore en avant ; un long poll a la suture, suivi de 4 autres plus courts, 
le dernier etant pre-antennal et tres court ; les 2 polls suturaux reunis par une ligne 
concave, suivie d'une ligne anguleuse qui aboutit aux bandes antennales ; celles-ci inco- 
lores, a bord interne sinueux. Fosse antennale peu profonde a son bord dorsal. Antennes 
fortes, incolores, sauf le sommet du 3 e article, qui est brunatre. l er article long, renfle, 
ovo'ide, une fois et demie aussi long que les 4 autres reunis ; 2 e ovo'ide et presque 
aussi long que les 3 suivants reunis ; le 3 e conique, recourbe en avant, plus long et 
plus gros que chacun des 2 suivants ; 4 e et 5" egaux, cylindriques, le 4 e insere sur la 
concavite du 3 e . Yeux saillants, au sommet d'une tache noire trapezoiidale. Bord 
temporal d'abord convexe et saillant, puis droit et dirige en dedans et en arriere ; 
un long poil a Tangle temporal. Bord occipital concave. Couleur generale blanchatre ; 
bandes incolores, les occipitales reduites a un espace triangulaire. 

Thorax long de mm. 9. Prothorax court, rectangulaire, pres de 2 fois aussi 
large que long, le bord posterieur un peu convexe, une epine a chaque angle posterieur ; 
blanchatre avec bande laterale incolore et recourbee un peu sur le bord posterieur. 
Me ta thorax un peu plus large que long, le plus large en arriere, a bords lateraux et 
posterieur concaves ; dans chaque angle posterieur 3 longs poils colores et inseres 
sur une me me saillie petite et conique ; une bande laterale courte, claire et saillante 
en dedans ; 3 sternites incolores (presternite lineaire, mesosternite en hexagone allonge 
transversalement, metasternite subcirculaire. Couleur generale blanchatre. Pattes 
incolores, sauf les tarses, qui sont brun chatain. 

Abdomen long de 2 mm. 3, plus large au 4 e segment. Segments a peu pres de 
memo longueur. A chaque angle posterieur, 2 longs poils au 1" segment ; 3 aux 
2 e , 3 e et 4 C ; 4 aux 5 e et 6" ; 6 au T ; 5 au 8 e ; le 9 e etroit, emargine a son extremite, 
dont chaque lobe porte un poil court et un poil long. Couleur generale blanchatre. 
Sur chaque segment, une bande laterale tres etroite, elargie en un prolongement 
transversal au bord anterieur et en un autre plus court en avant de I'angle posterieur ; 
au 7 segment les deux elargissements out la meme importance ; un seul au 8 e . Sur 
les segments 2 a 6 une petite tache noire dans Tangle anterieur ; une semblable au 8 e , 
suivie d'une tache marron ; chaque lobe du 9 C presque couvert par une tache marron ; 
Textremite de Tappareil copulateur est un peu coloree. Stigmates petits, situes sur le 
milieu de la longueur du segment, leur ecartement egal a la moitie de sa largeur. 



22 PROF. L. G. NEUMANN 

Femelle. Corps long de 4 mm. 2, large de 1 mm. Fosse antennale a angle anterieur 
formant une pointe aigue. Antennes greles ; l er article renfle, plus gros et presque 
aussi long que le 2 e ; l er et 2 e formant plus de la moitie de la longueur de 1'antenne ; 
3 et 5 e de meme longueur et plus longs que le 4". Abdomen long de 2 mm. 4 ; polls 
plus longs que cliez le $ ; bandes laterales ayant, sur chacun des segments 1 a 7, deux 
elargissements egaux. Pas de tache noire anterieure au 2 e segment, une au 7 e ; 8" 
segment plus etroit que chez le $ ; 9" a fente plus etroite. 

Cette espece est voisine de L. tricolor Piaget, et de L. confidens Kellogg. Elle 
rappelle aussi Nirmus giganticola Kellogg. 

5. Taschenbergius brevis Dufour 

1834. Philopterus brevis, L. Dufour, Ann. de la Soc. entomol. de France, iv., p. 674, 

PL XXL, fig. 3. 
1838. Philopterus (Docophorus ?) taunts Nitzsch ; H. Burmeister, Handbuch der 

Entomologie, ii., p. 433. 
1864. Docophoroides brevis, H. Giglioli, Quart. Journal of Microsc. Science (N.S.), iv., 

p. 21, PI. I., figs. 3 et 4. 

1866. Lipeurus taurus, C. G. Giebel, Zeitschrift f. ges. Naturwiss., xxviii., p. 385. 
1882. Eurymetopus taurus, 0. Taschenberg, Nova Acta der Ksl. Leop.-Carol.-Deutschen 

Akademie der Naturforscher, xliv., p. 183, PI. V., figs. 8 et 8a. 
1906. Taschenbergius brevis L. G. Neumann, Bull, de la Soc. Zoolog. de France, xx., 

p. 60. 

1 $ et 1 $, sur Diomedea exulans L., au S. de 1'Atlantique, par 35 lat. S. et 30 27' 
long. 0. ; 15 $ et 10 $, sur Diomedea exulans, au S. de 1'Ocean Indien, par 40 lat. S. et 
47 long. E. (avec Lipeurus hyalinus Nn.) ; 3 $ et 7 ?, sur D. exulans, dans 1'Ocean 
Indien, au S. de la Tasmanie (avec Lipeurus ferox Giebel, et L. hyalinus Nn.) ; 1 $ sur 
Thalassogeron culminatus Gould, au S. de 1'Ocean Indien, par 41 lat. S. et 53 24' 
long. E. 

T. brevis a encore pour hotes connus : Diomedea albatrus, D. nigripes, D. regia, 
Fulmarus glacialis, Puffinus opisthomelas, Arenaria interpres. 



PLATE III 



PLANCHE III 



TACIE 



FIGURE 1. Lipeurus hyalinus, n. sp 21 

a. $ Face dorsale x 20 

6. Abdomen de la ? x 20 

,, c. Antennes $ et $ . x 45 

,, d. Taches sternales du $ . x 65 

FIGURE 2. Philopterus lari, $, face dorsale 19 

FIGURE 3. Philopterus lari, $, tete et thorax (face ventralc) 
FIGURE 4. Philopterus lari, $, taches ventrales 



BRIT. ANTARCT. EXPED. 1907-9 



VOL. II. PLATE III 



NEUMANN: MALLOPHAGA 








3 





1. LlPEURUS HYALINUS, SP. N. 2-4. PHII.OPTE11CS tARI. 



G. Neumann del. 



BRITISH ANTARCTIC EXPEDITION 1907-9 

UNDER THE COMMAND OF SIR E. H. SHACKLETON, C.V.O. 

REPORTS ON THE SCIENTIFIC INVESTIGATIONS 

- 
VOL. II 

BIOLOGY 

EDITOR JAMES MURRAY 

PART IV 

A8TERIES, OPHIURES, ET ECHINIDES 

BY R. KOEHLER 

(LYONS) 



(WITH FIVE PLATES} 



LONDON 
PUBLISHED FOR THE EXPEDITION BY WILLIAM HEINEMANN 

21 BEDFORD STREET, VV.C. 

1911 

ISSUED SEPARATELY, JUNE, 1911 
PRICE 5*. NET 



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PART IV 

ECHINODERMA 

ASTERIES, OPHIURES, ET fiCHINIDES 

DE L'EXPEDITION ANTARCTIQUE ANGLAISE DE 1907-1909 

PAR R. KGEHLER 

Professeur a 1'UniversiW de Lyon 
(Avec les Planches IV & VIII) 

M. JAMES MURRAY a bien voulu m'offrir d'etudier les Asteries, Ophiures et Echinides 
recueillis par le Nimrod, commandant E. Shackleton, a la Terre Victoria du Sud, en 
1907-1909. Je suis heureux de le remercier ici de la confiance qu'il a bien voulu me 
temoigner. 

Les Echinoderm.es recueillis proviennent tous du Cap Royds, qui represente la 
pointe la plus occidentale de 1'ile Ross, dans la Terre Victoria du Sud, par 77 32' lat. S. 
et 166 12' long. E. La Terre Victoria et les regions avoisinantes ont deja etc explorees 
au point de vue zoologique ; la Southern Cross et la Discovery en ont rapporte des 
collections qui ont etc etudiees et plusieurs especes d'Echinodermes y sont connues. 
Aussi pouvait-on supposer que les Echinodermes du Nimrod n'offriraient pas un tres 
grand interet et ne renfermeraient que des representants d'une faune deja connue. 
Ces previsions n'ont pas ete realisees, heureusement : la collection qui m'a ete remise, 
bien qu'elle ne soit pas tres importante, est au contraire tres interessante, non pas 
seulement a cause des especes nouvelles que j'y ai trouvees, mais aussi en raison des 
renseignements qu'elle fournit sur la repartition geographique de certaines especes. 

Les explorations zoologiques de 1'Antarctique ont surtout ete faites dans deux 
directions differentes et presque opposees ; les unes, qui avaient pour point de depart 
le detroit de Magellan, ont eu pour objet 1'etude des regions situees au Sud de la 
pointe meridionale de 1'Amerique du Sad ; les Orcades et les Shetland du Sud, les 
terres de Graham, de Danco, etc., c'est-a-dire entre les 50 et 70 long. W. ; les autres^ 
qui avaient pour point de depart 1'Australie ou la Nouvelle-Zelande, nous ont fait 
connaitre des regions telles que la Terre Victoria du Sud, situees vers le 170 long. E. 
II reste entre ces deux points extremes un territoire a peine connu quoique la Belgica 
se soit avancee jusque vers le 100 long. W., et le Francais jusqu'au 126 long. W. 

BRIT. ANTARCT. EXPED. 1907-9. VOL. II. PART 4, ISSUED JUNE 1911 E 

B 



26 R. KCEHLER 

Les Echinodermes recueillis par le Nimrod comprennent six Asteries, quatre 
Ophiures, et trois Echinides, soit en tout treize especes dont huit sont nouvelles et 
trois representent des genres nouveaux. En voici 1'enumeration : 



ASTERIES 



Odontaster validus Kcehler. 
Cryaster antarcticus Koehler. 
Porania antarctica Smith. 



Ophioglypha resistens, n, sp. 
Ophioglypha flexibilis, n. sp. 



Sterechimis neumayeri Meissner. 



Coscinasterias brucei Koehler. 
Coscinasterias victories, n. sp. 
Notasterias armata, n. gen. u. sp. 



OPHIURES 



Amphiura algida, n. sp. 
Ophiodiplax disjuncta, n. gen. n. sp. 



ECHINIDES 



Pseudabatus nimrodi, n. gen. n. sp. 



Abatus shackletoni, n. sp. 

II est assez interessant de constater, qu'a part une seule et unique exception, 
aucune des especes ci-dessus n'a ete rapportee par la Discovery de la Terre Victoria 
du Sud ; la faune echinologique rencontree par ce batiment a une composition toute 
differente d'apres le rapport de J. Bell (08).* II me parait utile de rappeler ici les 
especes signalees par ce naturaliste : 



Asterias brandti Bell. 
Asterias longstaffi Bell. 
Heuresaster hodgsoni Bell. 
Pentagonaster incertus Bell. 
Leptoptychaster kerguelensis Smith. 
Cycethra verrucosa Philippi. 
Henricia ornata Perrier. 
Solaster octoradiatus Ludwig. 
Ophioglypha kcehleri Bell. 
Ophiozona inermis Bell. 
Ophiosteira antarctica Bell. 



Ophionotiis victor ice Bell. 
Ophiacantha imago Lyman. 
Ophiacantha vivipara Ljungmann. 
Ophiacantha cosinica Lyman. 
Ophioconis antarctica Lyman. 
Amphiura belgicce Koehler. 
Astrotoma agassizii I;yman. 
Jeune Ophiure. 

Austrocidaris canaliculata Agassiz. 
Sterechinus margaritaceus Lamarck. 
Abatus cavernosus Philippi. 



La seule espece de cette liste que j'ai retrouvee parmi les Echinodermes du Nimrod 
est la " jeune Ophiure " que J. Bell s'est contents de figurer sans la decrire ; nous 
verrons plus loin que cette forme, que le savant naturaliste anglais considerait comme 
un jeune, a tous les caracteres d'un adulte. 

Les especes deja connues que 1'Expedition Antarctique Anglaise a rapportees sont 
au nombre de cinq ; elles n'avaient pas encore ete signalees a la Terre Victoria. Deux 
d'entre elles sont plus ou moins communes dans le domaine antarctique ; ce sont la 
Porania antarctica et le Sterechimis neumayeri. La premiere espece est tres repandue 
dans les regions antarctiques et subantarctiques ; le Nimrod 1'a rencontree a une 
latitude tres elevee (77 -32' S.), alors que sa station la plus australe notee par la 
Belgica etait situee par 71 18'. Le Sterechinus neumayeri presente aussi une repartition 

Les chifl'res en caracteres gras renvoient i la liste des ouvrages cit6s qui se trouve 4 la fin du m6moire. 



ECHINODERMA 27 

geographique assez vaste : on le connait au detroit de Magellan, a la Georgia du Sud, 
aux Orcades du Sud, a la Terre de Graham, etc. ; Mortensen nous apprend (10, p. 68) 
qu'il a etc trouve en plusieurs stations par 1'Expedition Sud-polaire Suedoise et il estime 
qu'il doit etre circumpolaire ; sa repartition tient evidemment a ce qu'il possede une larve 
pelagique. II n'y a done rien d'etonnant a ce qu'il ait ete capture au Cap Royds. 

Les trois autres especes avaient jusqu'a present un habitat beaucoup plus restreint. 
La Coscinasterias briwei n'etait connue qu'aux Orcades du Sud ou la Scotia 1'a 
decouverte, et le Cryaster antarcticus a ete trouve par le Francais vers le 65 lat. S. 

Enfin VOdontaster validus a ete rencontre par le Francais vers les Terres de Graham 
et de Danco, et par la Scotia aux Orcades du Sud. Ces localites, comprises entre le 
50 et le 70 long. W., sont, on le voit, tres eloignees du Cap Royds. 

J'ai dit plus haut que les Echinodermes proviennent tous du Cap Royds. D'apres 
les renseignements qui m'ont ete communiques, les echantillons captures a une 
profondeur de 10 a 20 brasses ont ete recueillis dans une petite baie dont le fond 
renfermait une vase noire tres adherente ; ceux qui proviennent de 40 a 80 brasses 
ont ete dragues sur un fond de couleur claire dont la faune etait tres riche. 

ASTERIES 

ODONTASTER VALIDUS Kcehler 
Odontaster validus, Kcehler (06), p. 6 
Cap Royds. Juin 1908. Profondeur, 7-20 brasses. Quelques echantillons. 

L'un des exemplaires est de tres grande taille : R = G8 millim., r = 30 millim. ; 
il est tres robuste et 1'epaisseur au centre du disque depasse 20 millim. Cinq autres 
individus sont plus petits et R varie entre 38 et 25 millim. ; enfin d'autres individus, 
beaucoup plus petits et chez lesquels R est compris entre 15 et 8 millim., se trouvent 
dans la collection. 

PORANIA ANTARCTICA Smith 

Voir pour la bibliographic : 

Perrier (91), p. 107 et 163. 
Leitpold (95), p. 588. 
Ludwig (03), p. 22. 
Ludwig (05), p. 51. 

Cap Royds. 20 aout 1908. Profondeur, 60-80 brasses. Deux echantillons. 

Les exemplaires sont tres jeunes tous deux et ils ont la me me taille : R = 20 et 
r = 8 millim., mais dans 1'un d'eux le disque est plus petit tandis que les bras sont 
plus greles et plus longs que dans 1'autre. 

Les auteurs s'accordent a considerer comme devant rentrer dans la meme espece 
les P. antarctica Smith, magellanica Studer, glaber Sladen, et spinidata Sladen. 
Les Porania antarctiques varient en effet beaucoup dans leurs caracteres et Perrier 



28 R. KCEHLER 

a meme fait remarquer que certaines formes arrivaient a se rapprocher de la P. pulvillus 
des mers du Nord. 

Les deux individus que j'ai sous les yeux m'offrent un nouvel exemple de cette 
variation. L'un d'eux, qui a les bras plus courts et plus larges et le disque compara- 
tivement grand, offre, sur la face dorsale du disque, des tubercules coniques et bien 
apparents, au nombre d'une douzaine ; sur les bras, ces tubercules sont beaucoup 
plus petits, mais assez nombreux et rapproches. L'exemplaire correspond done, par 
ses caracteres, a la P.antarctica. Dans Fautre, dont les bras sont minces et allonges, 
il n'y a pas la moindre trace de tubercules ou de saillies sur la face dorsale : il 
rappellerait done la forme decrite par Sladen sous le nom de P. glaber. 

En raison des variations qui ont ete notees par differents auteurs sur les Porania 
des diverses regions antarctiques explorees, il ne semble pas qu'on puisse etablir une 
distinction entre les formes des iles Kerguelen et celles de 1'extremite meridionale de 
I'Amerique du Sud. 

La P. antarctica parait avoir une aire de repartition assez vaste dans les regions 
antarctiques. La Bdgica 1'a rencontree a 71 18' lat. S. : on voit qu'elle peut remonter 
encore davantage vers le pole puisque FEpedition Anglaise 1'a capturee a 77 32' 
lat. S. 

CRYASTER ANTARCTICUS Kcehler (PI. IV, fig. 1 et 2) 

Cryaster antarcticus, Koehler (06), p. 24, PI. I, fig. 1, et PI. II, fig. 10 
Baie de Backdoor. 6 fevrier 1907. Profondeur, 6-13 brasses. Un echantillon. 

L'exemplaire est en excellent etat de conservation. Au premier abord, il semble 
assez different du type du C. antarcticus que j'ai decrit d'apres les individus rapportes 
par la premiere Expedition du Dr. Charcot : tandis que cliez ces derniers les faces 
dorsale et ventrale du corps offrent des petits piquants nombreux et serres, 
Techantillon de 1'Expedition Antarctique Anglaise parait tout a fait inerme : de fait, 
ces piquants font a peu pres completement defaut et Ton ne trouve, en general, que 
la gaine tegumentaire des piquants, ceux-ci ayant disparu. Mais j'ai heureusement 
rencontre dans les collections recueillies recemment, au cours de la deuxieme Expedition 
du Dr. Charcot, et qui viennent de m'etre confies, des exemplaires de C. antarcticus 
chez lesquels les piquants sont beaucoup moins developpes que dans le type et peuvent 
rester completement caches dans leur enveloppe tegumentaire, de telle sorte que 
1'individu de la Terre Victoria apparait comme le dernier terme d'une serie dont je 
possede les principaux stades. II n'y a done aucune raison pour ne point le rapporter 
au C. antarcticus. 

Les bras, au nombre de cinq, sont subegaux, mais, comme ils sont plus ou moins 
contournes et releves Vers la face dorsale, il est difficile de mesurer leur longueur 
exacte : sur les plus grands bras, R varie de 120 a 130 millim. ; r = 49 millim. 

Le disque est tres epais, sa hauteur maxima atteignant 47 millim. ; il a la forme 
d'un cone surbaisse, a sommet tres emousse et il se continue largement avec les bras. 



ECHINODERMA 29 

On remarque, le long de chaque interradius, une depression assez large et qui 
s'accentue a mesure qu'on se rapproche de la peripherie du disque ; la base des bras 
se trouve ainsi nettement delimitee. Cette base est tres large : elle mesure 48 millim. 
sur le bras le plus large et 38 sur le plus etroit. Les bras se retrecissent tres rapide- 
ment sur leur premiere moitie, et ensuite d'une maniere plus lente, en s'amincissant 
progressivement jusqu'a 1'extremite qui est etroite et obtuse. 

Malgre 1'absence de depots calcaires dans les teguments de la face dorsale, celle-ei 
est tres resistante et indeformable et le tegument qui la recouvre est epais et dur. 
Tout 1'ensemble de 1'animal est tres robuste. La couclie superficielle de ce tegument, 
dont 1'epaisseur atbeint au moins un millimetre, est constitute par des saillies de forme 
tres irreguliere et inegales, qui sont tres serrees les unes centre les autres et separees 
par des sillons tres fins. Ces saillies out la surface tantot lisse, tantot plissee ; elles 
rappellent, par leur ensemble, les pustules que Ton observe sur la face dorsale de 
certaines Anasterias, mais elles sont beaucoup plus petites que dans ce dernier genre, 
car les plus grandes n'atteignent pas un centimetre de largeur. 

Les petits piquants que j'ai signales en 1906 chez les C. antarcticus provenant de 
la premiere Expedition Charcot, emergeaient de saillies analogues et leur base restait 
plongee, sur une longueur plus ou moins grande, dans le tissu mou qui lui formait 
ainsi une collerette. Ici le piquant fait completement defaut et la collerette du 
piquant existe seule. J'ai examine au microscope un grand nombre de ces pustules 
sans pouvoir y decouvrir la moindre indication de piquants ou de depots calcaires ; 
sur un certain nombre d'entre elles, on voit s'elever une petite protuberance centrale, 
conique, a pointe obtuse, mais celle-ci ne renferme pas non plus la moindre trace de 
piquant. On ne peut pas admettre que les piquants aient ete dissous par les reactifs 
conservateurs, car j'en observe sur la face ventrale ainsi que je le dirai plus loin. 

L'aspect de la face dorsale de cet individu est tout a fait identique a celui que 
j 'observe chez certains individus provenant de la deuxieme Expedition Charcot et 
chez lesquels cette face n'offre aucun piquant apparent ; mais 1'examen microscopique 
des collerettes ou pustules de ces individus montre que chacune d'elles renferme un 
petit piquant interne. 

Vers le centre du disque, les petites pustules s'aplatissent progressivement et elles 
disparaissent sur un espace mesurant 15 millim. de diametre environ, qui est occupe 
par un tegument lisse sur lequel on aperyoit cependant un fin reseau polygonal 
representant sans doute 1'impression de pustules disparues. II semble que cette 
region ait subi des pressions ou des frottements qui ont comprime ou enleve les parties 
superficielles du tegument. Au centre de cette region se trouve un tres petit pore 
qui correspond peut-etre a 1'anus. 

Parmi les pustules, se trouvent de tres nombreuses papules allongees et dont la 
longueur atteint 2 a 3 millim. ; leurs parois, tres molles, sont minces, transparentes 
et incolores et elles tranchent sur les autres parties des teguments dont la couleur 
est brun-clair. 



30 R. KCEHLER 

La couche ainsi constitute par les enveloppes des piquants avortes et les papules, 
atteint une epaisseur d'au moins 1 millim. ; en dessous d'elle se trouve une paroi 
conjonctive resistante, de couleur blanchatre, formee d'un tissu tres dense offrant 
des orifices circulaires par ou passent les papules. La couche superficielle des 
teguments donne, a la main, la sensation du velours. 

La plaque madreporique, tres grande et allongee dans le sens interradial, mesure 
10 millim. sur 7'5, et elle est situee plus pres du centre que du bord ; sa surface est 
grossierement granuleuse et les sillons ne sont pas apparents. 

Dans les aires interradiaires ventrales, les sillons qui separent les saillies tegu- 
mentaires se disposent d'une maniere reguliere, et parallelement les uns aux autres, 
en allant du sillon ambulacraire aux bords du disque et des bras. Les saillies tegu- 
mentaires elles-memes s'alignent ainsi en files transversales qui se continuent sur les 
bords du disque et des bras et passent aux pustules de la face dorsale. Ces saillies sont 
en general plus accentuees et plus grosses que celles de la face dorsale. Certaines d'entre 
elles, surtout au voisinage de la bouche, renferment un petit piquant que 1'on reconnait 
soit a 1'examen microscopique, soit en tatant 1'expansion a 1'aide d'une aiguille, mais 
aucun de ces piquants ne fait saillie au dehors. Sur les cotes des bras, mais dans la 
partie terminate seulement, j 'observe aussi quelques saillies coniques renfermant 
chacune un piquant interne. 

Les sillons ambulacraires sont etroits et les tubes sont places sans ordre, tres serres 
les uns contre les autres. Les piquants adambulacraires sont disposes sur trois rangs 
comme je 1'ai decrit ; ils sont tres courts, epais et entoures d'une forte enveloppe 
tegumentaire. 

COSCINASTEEIAS BRUCEI Kcehler (PI. V, fig. 5) 

Stolasterias brucei Koehler (08), p. 41, PI. V, fig. 46 et 47 
Cap Royds. 27 juillet 1908. Profondeur, 10-18 brasses. Quatre echantillons. 
Les quatre exemplaires sont tres jeunes et ils ont tous a peu pres la meme taille : 
R ne mesure pas plus de 27 millim. Ils n'ont pas encore acquis leur forme definitive, 
mais ils possedent tous les caracteres principaux de la C. brucei et je ne crois pas me 
tromper en les rapportant a cette espece que j'ai decrite d'apres les echantillons recueillis 
par la Scotia aux Orcades du Sud, a une profondeur de 10 brasses. Pour les raisons que 
j'indiquerai plus loin, il me parait plus correct de ranger cette Asterie dans le genre 
Coscinasterias plutot que dans le genre Stolasterias auquel je 1'avais d'abord rapportee. 
Les quatre individus presentent les memes caracteres : j'ai represente, PI. V, 
fig. 5, la face dorsale de 1'un d'eux que j'ai desseche pour rendre ses caracteres plus 
apparents. Le disque offre un cercle externe de piquants tres courts et obtus, entoures 
chacun d'une collerette a pedicellaires croises, peu nombreux, et, au centre, un petit 
piquant ou un groupe de deux ou trois piquants ; dans 1'espace intermediate, on 
n' observe que quelques piquants epars. La ligne carinale est marquee par une rangee 
de piquants qui offre, comrne chez le type de 1'espece, des sinuosites plus ou moins 



ECHINODERMA 31 

accentuees. Entre cette ligne et la rangee marginale dorsale, on n'observe que 
quelques piquants isoles, toujours entoures de leurs collerettes, mais la plus grande 
partie des' cotes des bras reste nue. Les piquants sont cylindriques et ils se terminent 
par une extremite arrondie offrant des denticulations tres fines et pointues. Les 
piquants des rangees marginales dorsales et ventrales sont aplatis et un peu plus gros 
que les precedents ; leur extremite est souvent un peu elargie et elle porte aussi de 
tres fines spinules. La rangee marginale ventrale est tres rapprochee des piquants 
adambulacraires dont elle est simplement separee par un intervalle etroit, sans la 
moindre trace de piquants. Ce caractere est bien conforme a ce qui existe chez la 
C. brucei, tandis qu'il existe une rangee ventrale de piquants chez la C. victories que 
je decrirai ci-dessous. 

Aux quatre echantillons que je viens de mentionner etaient joints une douzaine 
d'individus extreme me nt jeunes chez lesquels R est compris entre 12 et 4 millim. 
Ils appartiennent tres vraisemblablement aussi a la C. brucei. 

En rapportant primitivement la C. brucei au genre Stolasterias, j'avais donne a 
ce dernier genre la signification assez large indiquee par Sladen et non 1'acception 
restreinte qui lui a ete ensuite attribute par Perrier. Les differentes coupures qui 
ont ete etablies par ce dernier savant dans 1'ancien genre Stolasterias de Sladen ne 
sont peut-etre pas tres necessaires, et, d'autre part, les limites memes assignees par 
Perrier au genre Stolasterias ne sont pas bien precises. Ainsi ce naturalisle etablit en 
1896 (Stellerides des Campagnes de I'Hirondelle, p. 34), une distinction entre le genre 
Stolasterias et un nouveau genre Distolasterias qu'il cree, etqui est fonde sur le fait que le 
premier a les piquants adambulacraires isoles, tandis que le second en a deux par plaque. 
Or, trois pages plus loin (p. 37), il decrit, sous le norn de Stolasterias neglecta, une 
nouvelle espece qui possede deux rangees de piquants adambulacraires ; je ne puis com- 
prendre pourquoi 1'auteur ne la rapporte pas a son genre Distolasterias plutot qu'au 
genre Stolasterias qu'il caracterise, entre autres, par la rangee unique de piquants 
adambulacraires. 

D'autre part, W. K. Fisher a montre (06, p. 1104) que le genre Stolasterias cor- 
respondait exactement au genre Cosdnasterias, etabli par Verrill pour la C. muricata 
(qui est synonyme de C. calamaria Gray) ; le terme Cosdnasterias se trouve done 
anterieur au terme Stolasterias puisqu'il date de 1869, et le naturaliste americain 
estime que, logiquement, le premier doit etre substitue au second. Ce genre Cosdn- 
asterias, pris dans son sens le plus large, comprend les formes principales suivantes : 
calamaria, tenuispina, yemmifera, volsellata, stichantha, eustyla, gladalis, etc. 

Sans se prononcer d'une maniere formelle sur la necessite de subdiviser le genre 
Cosdnasterias, W. K. Fisher propose de substituer aux genres etablis par Perrier 
les coupures suivantes : 

Cosdnasterias Verrill, 1869 ; type, calamaria. 

Stolasterias Sladen, 1889 ( = Polyasterias Perrier); type, tenuispina. 



32 R. KCEHLER 

MartJiasterias Jullien, 1878 ; type, glatialis. 
Distolasterias Perrier, 1896 ; type, stichantha. 

II est bon de remarquer, a propos de ces quatre sous-genres, que le dernier n'a pas 
ete utilise par son createur dans le travail meme ou il 1'avait etabli, et que Favant- 
dernier a ete fonde sur un exemplaire mal conserve et n'offrant aucune valeur. 
D'autre part, 1'un des principaux caracteres sur lesquels on s'appuie pour separer 
des autres le genre Coscinasterias (comme aussi le genre Polyasterias de Perrier), est 
la multiplicite des bras. Dans ces conditions, il n'apparait pas d'une maniere bien 
evidente qu'il soit necessaire de subdiviser 1'ancien genre Coscinasterias de Verrill ; 
aussi j'ai cru devoir conserver ce terme en lui donnant sa signification la plus large 
sous laquelle il devient exactement synonyme du genre Stolasterias de Sladen. 

COSCINASTEBIAS VICTORLE, UOV. Sp. (PL V, fig. 3 et 4) 

Cap Royds. 20 aout 1908. Profondeur, 50-80 brasses. Un echantillon. 
Cap Royds. 30 aout 1908. Profondeur, 25 brasses. Un echantillon. 

L'exemplaire du 20 aout est en bon etat, bien qu'il ait ete legerement altere par un 
sejour dans le formol. Les bras, au nombre de cinq, sont un peu inegaux : dans le 
plus petit, R = 68 millim. ; chez d'autres, R varie de 75 a 82 millim. ; r 15 millim. 
Les bras sont tres legerement retrecis a la base et leur largeur maxima varie entre 
18 et 20 millim. ; ils se retrecissent progressivement jusqu'a 1'extremite qui est 
amincie et pointue. C'est cet individu qui me servira de type pour la description 
de 1'espece ; il est represente PI. V, fig. 3 et 4. 

L'individu capture le 30 aout n'est pas tres bien conserve ; il est quelque peu 
macere et ses tissus sont un peu ramollis ; le corps est aplati et il a du subir une 
certaine compression. Mesure sur le bras le plus grand, R depasse 130 millim. ; les 
bras atteignent 27 millim. de largeur a la base et ils s'amincissent progressivement 
jusqu'a 1'extremite qui est pointue ; r = 20 millim. 

Le disque est de taille moyenne : son diametre, mesure entre deux espaces inter- 
brachiaux non consecutifs, oscille autour de 30 millim. La face dorsale des bras est 
convexe, legerement carenee sur la ligne mediane ; les faces laterales, comprises entre 
les rangees de plaques marginales dorsales et ventrales, sont etroites. Le squelette 
des bras est assez resistant et 1'ensemble de 1'animal est assez rigide. 

La face dorsale du disque et des bras offre des piquants peu developpes qui 
emergent d'une collerette ressemblant plutot a une pustule d'Anasterias ; ces 
collerettes sont accolees les unes centre les autres, et, entre elles, se montrent de 
nombreuses papules saillantes, de telle sorte que le squelette sous-jacent est absolu- 
ment invisible. Les collerettes, avec les piquants qu'elles enveloppent, sont beaucoup 
moins distinctes que dans la C. brucei et Ton reconnait moins nettement que dans 
cette derniere espece les rangees qu'elles forment. 

La region centrale du disque, sur un diametre de 14 millim., est occupee par un 



KCIIINODKKMA 33 

premier cercle de piquants entoures de leurs collerettes qui sont contigues, inegales 
et peu distinctes : ces piquants font a peine saillie hors de la collerette et leur 
extremite se termine en pointe emoussee. En dedans, on reconnait un deuxieme cercle 
de piquants moins apparents, et dont les collerettes sont encore plus petites et moins 
distinctes que les precedentes. Le reste de 1'espace circulaire est occupe par des 
collerettes depourvues de piquants, sauf au centre ou il existe un piquant un peu plus 
fort que les autres. Entre les collerettes se montrent de nombreuses papules. 

La ligne carinale des bras offre une rangee de piquants qui partent du cercle 
exterieur signale ci-dessus et qui depassent a peine la collerette entourant leur base. 
Ces collerettes sont serrees les unes contre les autres et leurs limites ne sont pas tres 
apparentes en general. Elles forment une ligne irregulierement sinueuse, et, si leurs 
contours ne sont pas tres distincts, la rangee qu'elles constituent sur chaque bras est, 
dans son ensemble, assez reconnaissable en general ; sur la photographic, ces rangees 
ne sont pas tres apparentes. Les dimensions de ces collerettes decroissent tres lente- 
ment, mais elles deviennent tres petites dans le dernier quart des bras, et, par con- 
sequent, elles s'y montrent tres nombreuses ; leur piquant central reste distinct 
jusqu'a 1'extremite du bras. Je compte plus de soixante collerettes dans cette rangee 
carinale : elles sont, comme on le voit, plus petites, plus nombreuses, et plus serrees 
que chez la C. brucei ou leur nombre ne depasse pas quarante-cinq dans un exemplaire 
de meme taille. On peut constater, en comparant la photographic que je donne ici 
(PI. V, fig. 3) de la C. victories au dessin que j'ai public de la C. brucei (08, PI. V, fig. 46), 
que, dans cette derniere, les collerettes conservent les memes dimensions sur presque 
toute la longueur des bras et qu'elles sont a peine plus rapprochees et plus petites 
vers 1'extremite, tandis qu'ici il en est tout autrement. Les sinuosites sont plus 
accentuees chez la C. victories, mais la ligne elle-meme est, dans son ensemble, moins 
distincte que chez la C. brucei. L'espace, tres large, compris entre la rangee carinale 
et la rangee marginale dorsale, est occupe par des collerettes confluentes dont les 
contours sont presque toujours mal indiques, et qui sont un peu plus petites et moins 
developpees que celles de la rangee carinale ; on reconnait, par endroits, un arrange- 
ment en rangees longitudinales au nombre de deux ou trois de chaque cote. Du 
centre de chaque collerette, s'eleve habituellement un petit piquant tres court et 
obtus. Enfin, entre les collerettes, se montrent de nombreuses papules. 

Sur le grand exemplaire, les contours des collerettes sont mieux marques : elles 
sont arrondies ou irregulierement polygonales par pression reciproque. Celles de la 
ligne carinale ne sont pas beaucoup plus grandes que les autres et elles mesurent 
environ 2'5 millim. de largeur ; elles sont disposees suivant une ligne beaucoup moins 
sinueuse que sur le premier individu, mais plus distincte en revanche. Le piquant 
central reste toujours peu saillant. Les collerettes laterales forment trois ou quatre 
rangees longitudinales assez regulieres et elles constituent egalement des rangees 
transversales legerement obliques plus ou moins apparentes, dans chacune desquelles 
elles sont le plus souvent au nombre de quatre. La forme de ces collerettes est 

BRIT. ANTARCT. EXPED. 1907-9. VOL. II. P 



34 R. K(EHLER 

irregulierement polygonale, et leur largeur atteint 2 millim. Elles sont bien separees 
les unes des autres, aussi distinctes et presque aussi grosses que celles de la rangee 
carinale et leurs limites sont beaucoup mieux marquees que dans 1'autre exemplaire. 
Chacune d'elles porte un piquant central, court et emousse. Les papules sont toujours 
abondantes et elles sont surtout nombreuses vers les bords des bras ou elles forment 
de petits groupes reguliers entre les collerettes successives. 

Le bord dorsal des bras est occupe par une rangee de gros piquants aplatis, courts, 
a extremite tronquee et legerement elargie, qui s'elevent chacun du centre d'une 
collerette ; leur region libre mesure environ 1'5 millim. de hauteur. La surface de 
ces piquants est legerement canaliculee. Les collerettes, de forme rectangulaire, sont 
un peu plus grosses que celles de la face dorsale des bras. Vers 1'extremite des bras, 
les piquants sont plus petits et plus serres ; ils deviennent alors tres nombreux, ainsi 
que cela arrive pour la rangee carinale. 

Les piquants de la rangee marginale ventrale correspondent a ceux de la rangee 
marginale dorsale, mais ils sont plus epais et plus forts, tout en conservant la meme 
forme generale. Les collerettes qui en entourent la base sont identiques a celles de 
la rangee dorsale. Entre ces deux rangees de piquants marginaux, les faces laterales 
des bras, tres etroites, sont occupees par des papules tres serrees et meme tout a fait 
contigues, formant une bande dont la largeur, a la base des bras, ne depasse pas deux 
millimetres. 

La face ventrale des bras presente une rangee de piquants qui correspondent a 
peu pres exactement a ceux de la rangee marginale ventrale, mais qui sont un peu 
moins developpes qu'eux. Ils deviennent plus petits dans la seconde moitie des bras, 
et, dans le dernier quart, ils ne se montrent qu'a des intervalles assez espaces. Cette 
rangee est tout a fait contigue aux piquants adambulacraires et a ceux de la rangee 
marginale ventrale sur I'exemplaire qui me sert de type ; sur le plus grand, elle est 
separee des premiers par un espace etroit mais cependant bien apparent : mal- 
heureusement la face ventrale de ce dernier individu est en si mauvais etat qu'elle ne 
se prete pas a 1' etude. 

Les sillons ambulacraires sont tres larges et les tubes forment quatre rangees 
irregulieres. Les piquants adambulacraires sont disposes sur deux rangs : ils sont 
assez courts, mais fortement aplatis dans le sens transversal ; leur extremite est tronquee 
et arrondie ; les deux rangees sont a peu pres identiques. Cinq piquants de la rangee 
externe correspondent a peu pres a deux piquants de la rangee marginale ventrale. 

Tous ces piquants, comme d'ailleurs ceux des rangees ventrales et marginale s 
ventrales, sont enveloppes d'une enveloppe tegumentaire assez epaisse qui les recouvre 
completement. 

Les dents portent, a leur pointe proximale, chacune deux piquants cylindriques, 
termines par une extremite arrondie et plus minces que les adambulacraires ; sur 
leur face ventrale, on rencontre un piquant plus fort, avec 1'extremite tronquee et 
rappelant davantage les piquants adambulacraires dont il differe cependant par sa 
forme cylindrique. 



ECHINODERMA 35 

II existe, comme d'habitude, des pedicellaires croises assez nombreux dans le tissu 
des collerettes, et des pedicellaires droits qui se montrent, de distance en distance, sur les 
parois du sillon ambulacraire. Malheureusement le tissu calcaire de ces petits organes 
a ete quelque pen attaque par le forinol et Ton ne peut en reconnaitre la structure. 

Rapports et differences. La C. victories est tres voisine de la C. bnicei ; elle s'en 
distingue par ses bras plus amincis et comparaiivement plus longs, par le nombre 
des piquants carinaux, beaucoup plus eleve parce que ces piquants deviennent tr&s 
petits et serres dans la partie terminate des bras, par les piquants plus nombreux 
sur la face dorsale des bras ou ils forment meme plusieurs rangees longitudinales 
assez reconnaissables dans le grand exemplaire et enfin par la presence d'une rangee 
distincte de piquants sur la face ventrale, entre les piquants adambulacraires externes 
et la rangee marginale ventrale. Les piquants des deux rangees adambulacraires, 
ainsi que ceux des rangees ventrales et marginales ventrales, sont enveloppes d'une 
enveloppe tegumentaire qui les recouvre completement, disposition que je n'ai pas 
observee chez la C, brucei. 



NOTASTEKIAS, nov. gen. 

Le squelette des bras comprend cinq rangees regulieres de plaques, une carinale, 
deux marginales dorsales et deux marginales ventrales, ainsi que cela est la regie 
dans le genre Coscinasterias et les genres voisins. II existe en outre un commence- 
ment de rangee ventrale. Chaque plaque porte un piquant unique assez developpe, 
mais depourvu de collerettes a pedicellaires croises. Les piquants de la rangee carinale 
offrent, a leur base, chacun un pedicellaire a valves croisees mais appartenant a un 
type particulier, et dont les dimensions sont vraiment considerables puisque leur 
longueur peut atteindre et meme depasser 2 millim. Ces pedicellaires sont toujours 
isoles a la base du piquant ; ils ont les memes caracteres que ceux que j'ai decrits 
chez YAsterias pedicellaris recueilli par la Scotia (08, p. 49) ; leur structure differe 
de celle des pedicellaires forcipiformes ordinaires et je propose de leur donner le nom 
de macrocepji.ales ; je reviendrai plus loin sur leurs caracteres. A la base des piquants 
portes par les plaques marginales dorsales et ventrales, on rencontre aussi un et 
parfois deux pedicellaires macrocephales, mais ces pedicellaires sont beaucoup plus 
petits que ceux qui accompagnent les piquants carinaux. Certains de ces piquants 
marginaux portent en outre des pedicellaires croises ordinaires, au nombre d'un ou 
de deux au plus, et s'inserant sur le piquant lui-meme entre sa base et son milieu. 
Ces pedicellaires forcipiformes sont toujours plus petits que les plus petits pedicellaires 
macrocephales voisins : ils mesurent en effet 0'4 millim. de longueur environ, tandis 
que les plus petits pedicellaires macrocephales atteignent pres du double. 

Dans la seule espece connue, qui est de taille plutot petite, il n'y a pas de rangee 
de plaques dorso-laterales. Les papules sont isolees et bien distinctes, arrondies et 



36 R. KtEHLER 

relativement assez grandes ; elles sont espacees et forment une premiere rangee entre 
les carinales et les marginales dorsales, puis une deuxieme, plus reguliere, entre les 
marginales dorsales et ventrales. Les piquants adambulacraires sont disposes suivant 
quatre rangees irregulieres. II existe des pedicellaires droits qui se montrent de 
distance en distance sur les parois des sillons ambulacraires. Les tubes ambulacraires 
forment plusieurs series irregulieres. 

II me parait utile de revenir sur les caracteres des pedicellaires macrocephales que 
j'ai pu etudier sur les deux echantillons de 1' Expedition Anglaise, ainsi que sur un 
troisieme exemplaire provenant de la deuxieme Expedition Charcot, d'une maniere 
plus complete que chez YAsterias pedicettaris ou ces gros pedicellaires etaient assez rares. 
Cette etude m'a convaincu que les differences entre ces pedicellaires et les pedicellaires 
forcipiformes sont assez marquees pour que Ton doive considerer les premiers comme 
representant une forme a part ; c'est pourquoi j'ai cru devoir leur appliquer un nom 
particulier. 

Exterieurement, les pedicellaires macrocephales se reconnaissent non seulement 
a leur taille, mais aussi a leur forme qui est conique, les deux valves qui constituent 
chacun d'eux allant en se retrecissant jusqu'a I'extremite, au lieu de former une lame 
convexe dont le bord libre est large, arrondi et muni d'une serie de fines denticulations. 
En outre, I'extremite de chaque valve est recourbee et se termine par un crochet plus 
ou moins developpe qui se croise avec son congenere de telle sorte qu'on pourrait dire 
que ces pedicellaires sont doublement croises : en effet, leurs valves se croisent d'abord 
au niveau de leur articulation sur la piece basilaire, puis, une deuxieme fois, vers leur 
extremite. On peut voir, en comparant les dessins que je donne ici des pedicellaires 
macrocephales (PI. V, fig. 6 a 11 ; PI. VI, fig. 4 a 8) aux figures de pedicellaires 
forcipiformes qui ont ete publiees, soit autrefois par Perrier, soit plus recernment par 
Ludwig (03, PI. VII, fig. 66 a 68), combien les deux formes sont differentes. Dans les 
gros pedicellaires d'Asterias pedicettaris, le crochet terminal n'etait pas tres developpe, 
mais il etait toujours tres fort ettres apparent sur les petits (voir Koehler, 08, PL VIII, 
fig- 75). 

Ainsi que je 1'ai dit en 1908, les valves de nos pedicellaires sont creuses : elles ont 
la forme d'un cornet dont une partie du bord libre se continue en une " queue " qui 
sert a Farticulation avec la piece basilaire. En d'autres termes, la lame calcaire qui 
constitue la valve, s'enroule sur elle-meme sur la moitie de sa longueur environ, et 
les deux bords de la partie enroulee s'adossent 1'un a 1'autre suivant une ligne droite 
qui porte des dents coniques et pointues (PL V, fig. 8, 9 et 11 ; PL VI, fig. 5 et 6). 
Ces dents sont moins nombreuses et moins fortes sur les gros pedicellaires que sur les 
petits, et elles ne se montrent que sur la partie proximale de la ligne suturale, de telle 
sorte que toute la region qui precede le crochet est depourvue de dents. Sur les petits 
pedicellaires, les dents sont plus nombreuses et elles se montrent sur une plus grande 
longueur le long de la ligne suturale ; chez YA. pedicettaris, elles peuvent meme 
s'avancer jusqu'au voisinage du crochet. Elles s'engrenent avec leurs congeneres de 



ECHINODERMA 37 

la valve opposee. Le crochet terminal est pointu, plus ou moins allonge, tantot droit, 
tantot legerement recourbe. 

La queue de la valve, assez longue, est elargie, concave et elle se termine par un 
bord arrondi. 

La lame calcaire qui forme chaque valve est constitute par un tissu areole, avec 
des perforations petites et tres serrees, et la meme structure s'observe sur toute 1'etendue 
de la queue qui n'est pas constitute par du tissu compact. Seul le crochet qui termine 
les valves est forme par un tissu compact et transparent. 

On voit done que les valves des pedicellaires macrocephales, avec leur forme de 
cone ou de cornet, leur crochet terminal, leurs denticulations disposees le long de la 
ligne suturale qui correspond a 1'une des generatrices du cone, et la queue formee de 
tissu calcaire areole, sont bien differentes de celles des pedicellaires croises ou forcipi- 
formes. La piece basilaire sur laquelle les valves s'articulent, offre au contraire une 
composition peu differente de celle que Ton connait chez ces derniers (PI. V, fig. 10, et 
PI. VI, fig. 7). Son corps represente une lame aplatie, allongee dans le genre 
Notasterias, plus courte chez V Asterias ped-icellaris, et qui se termine par un bord 
fortement convexe : celle-ci se presente par sa tranche quand on regarde le pedi- 
cellaire de profil comme celui qui est represente PI. V, fig. 6 et 7, tandis que lorsque le 
pedicellaire est vu de face (PI. VI, fig. 4 et 8), la lame se montre a plat. Les perfora- 
tions, petites et nombreuses, sont disposees en rangees lineaires, legerement divergentes 
et extremement serrees. La base de cette lame s'epaissit sur ses cotes de maniere 
a former deux bords arrondis qui s'etendent presque perpendiculairement a son plan et 
se continuent chacun en avant et en arriere par une sorte d'apophyse conique et arrondie ; 
ces apophyses correspondent respectivement aux apophyses regulieres et irregulieres de 
Perrier. On remarque que 1'une des apophyses d'un cote est plus forte que 1'autre 
du meme cote, mais il n'y a pas, dans le developpement et la direction, une inegalite 
comparable a celle que 1'on connait chez les pedicellaires forcipiformes. Dans les 
petits pedicellaires macrocephales, les apophyses sont plus pointues et plus allongees 
et la lame est au contraire plus courte ; aussi, quand on regarde la piece basilaire par 
le cote, on voit sur le plan superieur les deux apophyses d'un cote et sur le plan 
inferieur les deux autres apophyses, tandis que la piece basilaire ne represente qu'une 
saillie peu marquee. C'est 1'aspect que j'ai figure chez YA. pedicellaris (08, PI. VII, 
fig. 67) : la lame, moins developpee que dans le genre Notasterias, etait tout a fait 
invisible. 

Les muscles adducteurs des valves sont extremement developpes ; on les apergoit 
par transparence sur des pedicellaires montes en entier et ils se prolongent assez haut 
dans 1'interieur des valves (PI. V, fig. 7, et PL VI, fig. 4). 

Les pedicellaires macrocephales sont toujours isoles ; ils ne sont jamais reunis en 
collerettes a la base des piquants comme Ton observe dans les Asterias et les genres voisins. 
Leurs valves sont entourees par un tissu conjonctif transparent qui se continue, a la 
base du pedicellaire, par un pedicule extremement court et mince, a 1'aide duquel le 



38 R. KCEHLER 

pedicellaire se rattache au tegument de 1'Asterie. II n'y a pas de cordon fibreux 
s'inserant sur la piece basilaire comme dans les pedicellaires forcipiformes. 

La presence d'une forme particuliere de pedicellaires chez une Asterie m'a paru 
suffisante pour justifier la creation d'un genre nouveau, d'autant plus que les pedi- 
cellaires droits ou croises conservent chez toutes les Asteriadees une structure tres 
uniforme. Je propose d'appliquer le nom de Notasterias a 1' Asterie decouverte par 
1'Expedition Antarctique Anglaise. 

Ainsi que j'ai eu Foccasion de le rappeler plus haut, j'ai rencontre les memes pedi- 
cellaires macrocephales chez un Asterie recueillie par 1'Expedition Antarctique 
E'cossaise, a une profondeur de 1410 brasses, et que j'ai decrite sous le nom d'Asterias 
pedicellaris, mais cette derniere ne saurait rentrer dans le genre Notasterias : elle en 
diflere, en efEet, par la constitution de son squelette, par la repartition irreguliere des 
pedicellaires macrocephales, par 1'absence de pedicellaires forcipiformes et par les 
piquants adambulacraires ne formant qu'une seule rangee. Je crois qu'il est necessaire 
d'enlever cette espece au genre Asterias et je serais dispose a en faire le type d'un 
nouveau genre qu'on pourrait appeler Aittasterias. Les caracteres distinctifs de ces 
deux genres antarctiques peuvent se resumer de la facon suivante : 

Notasterias. Le squelette des bras est forme par cinq rangees longitudinales de 
plaques assez grandes et unies de maniere a former un squelette compact ne laissant 
que des orifices petits pour le passage des papules qui sont isolees. Chaque plaque 
porte un piquant depourvu de collerettes a pedicellaires forcipiformes. Les piquants 
carinaux offrent, a leur base, chacun un gros pedicellaire macrocephale unique ; des 
pedicellaires analogues, mais plus petits, peuvent se rencontrer a la base des piquants 
marginaux dorsaux et ventraux, et ces piquants peuvent en outre porter quelques 
pedicellaires forcipiformes. Les piquants adambulacraires sont disposes sur deux 
rangees. 

Une seule espece, antarctique et littorale. 

Autasterias. Le squelette des bras est forme de cinq rangees longitudinales de 
plaques petites, portant chacune un piquant et reliees par des travees qui laissent 
entre elles de tres larges mailles. Sur le reseau calcaire de la face dorsale, mais non 
specialement a la base de chaque piquant, se trouvent dissemines quelques gros 
pedicellaires macrocephales. II n'y a pas de pedicellaires a la base des piquants 
portes par les plaques carinales et marginales dorsales, mais, a la base de chaque 
piquant marginal ventral, il existe un groupe de quelques petits macrocephales ; les 
pedicellaires forcipiformes font completement defaut. Les piquants adambulacraires 
ne forment qu'une seule rangee. 

Une seule espece, antarctique et abyssale. 



ECHINODERMA 39 

NOTASTEEIAS ARMATA, UOV. Sp. (PL V, fig. 6 a 11 ; PI. VI, fig. 1 a 8) 

Bale du Cap Royds. 2 juillet 1908. Profondeur, 10-18 brasses. 

Deux echantillons. 

Les bras sont un pen inegaux. Dans le plus grand individu, R = 25 a 28 millim., 
r = 5'5 millim. Dans le second, R = 11 a, 20, r = 4 millim. 

Le tegument est assez epais et il cache coinpletement les contours des plaques 
sous-jacentes ; arm de pouvoir etudier ces dernieres, j'ai desseche le plus petit 
echantillon chez lequel les contours des plaques sont devenus bien apparents. Je 
represente ici le plus grand individu (PL VI, fig. 1 et 2). 

Le disque est petit ; les bras en sont bien distincts, mais ils ne sont pas retrecis a 
la base. Leur largeur est de 6'5 millim. en moyenne ; elle ne diminue guere que dans 
le dernier quart et Fextremite est amincie. 

La face dorsale du disque offre, a sa peripherie, un cercle de plaques au nombre de 
dix dans le petit exemplaire, cinq radiales et cinq interradiales ; la disposition parait 
etre la meme dans le grand exemplaire. Le centre est occupe par une plaque plus 
grande qui se relie a celles de la peripherie par des ossicules rayonnants. Chaque 
plaque porte un piquant assez fort, mesurant 2'5 a 3 millim. de longueur, assez elargi 
a la base et devenant ensuite cylindrique ; Fextremite, obtuse, offre des rugosites ou 
de fines denticulations ; de meme que les piquants des bras, ceux-ci sont recouverts 
d'une mince enveloppe tegumentaire. A la base de chaque piquant se trouve un 
gros pedicellaire macrocephale identique a ceux que nous retfouverons sur les bras, 
mais un peu plus petit en general que ces derniers, sauf celui qui se trouve a la base 
du piquant central. La plaque madreparique est petite, allongee dans le sens inter- 
radial et placee a peu pres a egale distance du centre et des bords ; ses sillons sont 
peu marques. 

Les bras presentent cinq rangees longitudinales de plaques : une carinale, deux 
marginales dorsales et deux marginales ventrales. La rangee carinale est bien 
saillante, de telle sorte que les bras sont assez fortement carenes ; la rangee marginale 
dorsale est aussi tres accusee. La rangee marginale ventrale est assez eloignee de la 
precedente et separee d'elle par une face verticale ; la coupe du bras se rapproche 
ainsi d'un polygone presque regulier, et dont la base seule est un peu plus grande que 
les autres cotes. Les plaques carinales, fortes et saillantes, ont une forme trifoliee, 
due a la presence de trois lobes arrondis, le lobe proximal recouvrant la region mediane 
de la plaque precedente et les lobes lateraux se reliant par de petites rangees aux 
plaques marginales dorsales. Les trois ou quatre premieres plaques de chaque rangee 
portent chacune un piquant, mais les suivantes ne le possedent pas toujours et parfois 
on ne le rencontre qu'une fois sur deux. II y a une dizaine de piquants sur chaque 
bras dans le grand exemplaire et huit en moyenne dans le petit. Les plaques qui 
portent un piquant sont plus fortes que les autres. Ces piquants ont les memes 
caracteres que ceux du disque ; ils sont seulement un peu plus longs et leur longueur 



40 R. K(EHLER 

atteint generalement 3 inillim. Us sont absolument depourvus de collerettes a 
pedicellaires, mais, a la base de cnacun d'eux, se trouve un gros pedicellaire 
macrocephale dont la longueur peut atteindre pres de 3 millim. En raison de 
ses dimensions, ce pedicellaire rejette souvent le piquant, soit d'un cote, soit de 
1'autre ; comme, d'autre part, la taille de ces pedicellaires varie, il en resulte que la 
rangee carinale de piquants parait plus ou moins irreguliere. 

Les plaques marginales dorsales ont la meme forme et la meme disposition que les 
carinales, mais elles sont un peu plus saillantes que ces dernieres ; leurs lobes lateraux 
internes ne se reunissent directement a ceux des plaques marginales correspondantes 
qu'a 1'extremite des bras, mais, sur le reste de la longueur des bras, on remarque deux 
ou trois petites plaques intercalaires. La plupart des marginales dorsales portent 
un piquant rappelant ceux des carinales, mais un peu plus petit, et ces piquants 
paraissent plus frequents que sur ces dernieres plaques. En principe, les plaques 
marginales dorsales correspondent aux carinales, mais la correspondance est parfois 
troublee en raison de la presence ou de 1'absence de piquants sur les plaques con- 
siderees. Certains piquants du commencement des bras offrent, a leur base, un 
pedicellaire macrocephale constitue comme ceux de la serie carinale, mais un peu 
plus petit ; ces pedicellaires macrocephales ne sont pas tres nombreux sur les deux 
echantillons de 1'Expedition Anglaise ; j'aurai 1'occasion de faire remarquer plus loin 
que dans un exemplaire de Notasterias armata, recueilli par la deuxieme Expedition 
Charcot, ces pedicellaires macrocephales sont assez nombreux sur la serie marginale 
dorsale. Ici, ces pedicellaires ne se montrent que sur les premiers piquants de la 
rangee marginale dorsale et les suivants portent, en general, chacun un ou deux 
pedicellaires forcipiformes ordinaires, qui s'inserent, non plus a la base du piquant, 
mais a une hauteur variable sur sa premiere moitie, comme on 1'observe chez diverses 
Asterias. Quand il y a deux pedicellaires croises sur le meme piquant, ceux-ci s'inserent 
au meme niveau ; ils peuvent d'ailleurs coexister avec un pedicellaire macrocephale 
place a la base du piquant. Ainsi que je 1'ai dit plus haut, ces pedicellaires croises 
sont toujours plus petits que les plus petits pedicellaires macrocephales. 

Les plaques marginales ventrales sont encore plus saillantes que les marginales 
dorsales et elles forment, par leui ensemble, un bord tranchant qui separe la face 
ventrale du reste du corps. Elles correspondent exactement aux marginales dorsales 
dont elles sont assez eloignees et auxquelles elles sont reunies par des arceaux tres 
reguliers, disposes parallelement et comprenant chacun quelques petites plaques : 
ainsi se trouvent determinees les faces laterales qui sont verticales. Chaque plaque 
porte un piquant analogue a ceux des marginales dorsales, mais un peu plus petit, 
legerement aplati, avec 1'extremite arrondie et munie de fines asperites. La plupart 
de ces piquants presentent, a leur base, un pedicellaire macrocephale plus petit que 
ceux de la rangee carinale, et, plus haut, un ou deux petits pedicellaires croises 
ordinaires s'inserant sur le piquant lui-meme. 

Entre les petits arceaux de plaques qui relient les carinales aux marginales dorsales, 



ECHINODERMA 41 

on remarque une ligne reguliere de papules arrondies, absomment isolees et assez 
grandes ; on pent me me observer, a la base des bras, le commencement d'une deuxieme 
rangee. Sur les cotes verticaux des bras, entre les plaques marginales dorsales et 
ventrales, il existe une autre rangee, tres reguliere, de papules analogues. 

Les sillons ambulacraires sont larges et renferment quatre series irregulieres de 
tubes serres. 

Les piquants adambulacraires sont disposes suivant deux rangees tres regulieres : 
ces piquants sont cylindriques ou aplatis par suite de leur pressionreciproque, et leur 
extremite est arrondie ; ils sont couverts d'une mince enveloppe tegumentaire qui 
les fait paraitre lisses, mais, quand ils sont desseches, on constate qu'ils sont couverts 
d'asperites extremement fines et rapprochees. 

Sur les parois du sillon, on reconnait, de distance en distance, des pedicellaires 
droits qui ne presentent rien de particulier, et dont la longueur moyenne est d'un 
millimetre (PI. VI, fig. 3). 

Sur le grand exemplaire, il existe a la base des bras, entre les piquants adambula- 
craires et la rangee des piquants marginaux ventraux, une rangee intercalaire de 
piquants plus petits que ces derniers et qui leur correspondent assez exactement. 
Cette rangee ventrale atteint un developpement variable : tantot elle disparait avant 
le milieu du bras, tantot elle s'etend un pen plus loin. Ces piquants sont en general 
depourvus de pedicellaires : j'observe cependant un pedicellaire croise sur deux d'entre 
eux. Dans le petit exemplaire, cette rangee de piquants fait a pen pres completement 
defaut : on en retrouve cependant quelques vestiges a la base des bras. 

Parmi les Echinodermes qui ont ete recueillis par la deuxieme Expedition 
Antarctique Francaise du Dr. Charcot, et qui viennent de m'etre confies, se trouve 
un exemplaire de Notasterias armata qui porte a trois le nombre des individus actuelle- 
ment connus de cette espece. Les dimensions sont voisines de celles des deux 
echantillons de la Terre Victoria : R - 20 a 25 millim. Les deux rangees de plaques 
marginales dorsales sont un pen plus rapprochees de la rangee carinale et les gros 
pedicellaires macrocephales sont un peu plus abondants : ils se montrent d'une maniere 
assez constante a la base des piquants de la rangee marginale dorsale et j'en retrouve 
meme a la base d'un certain nombre de piquants marginaux ventraux, mais ils sont 
alors beaucoup plus petits. En revanche, les pedicellaires croises ordinaircs sont 
tres rares : j'en observe cependant quelques-uns sur les piquants des plaques marginales 
ventrales. 



BRIT. ANTARCT. EXPED. 1907-9. VOL. II. G 



42 R. KCEHLER 



OPHIURES 

OPHIOGLYPHA RESISTENS, nov. sp. (PI. VII, fig. 9 a 12) 
Cap Royds. Profondeur 10-20 brasses. l er juillet 1908. Plusieurs echantillons. 

Dans le plus grand exemplaire, le diametre du disque est de 12 millim. et les bras 
n'ont que 21 a 22 millim. de longueur : en general, les bras sont un peu plus longs et 
dans un echantillon chez lequel le diametre du disque est de 10 millim. seulement, 
leur longueur atteint 28 millim. L'ensemble est tres robuste : le disque est epais ; 
les bras sont forts, epais et carenes sur la ligne mediane dorsale. 

Le disque est arrondi ou subpentagonal. La face dorsale est couverte de plaques 
assez nombreuses, tres inegales, epaisses et separees par de larges sillons. On distingue 
generalement une centro-dorsale, grande et arrondie, et, en dehors, un cercle de cinq 
radiales un peu plus petites qu'elle et elargies transversalement : ces plaques sont 
separees les unes des autres par une rangee de deux petites plaques arrondies; elles 
sont egalement separees de la centro-dorsale par un cercle de plaques plus petites, 
entre lesquelles se montrent d'autres plaques tres reduites. En dehors de cette partie 
centrale, viennent d'autres plaques parmi lesquelles on remarque, dans les espaces 
radiaux, une assez grande plaque arrondie, situee a la base de chaque paire de 
boucliers radiaux et a laquelle fait suite une rangee de deux ou trois petites plaques 
separant ces boucliers radiaux 1'un de 1'autre. Dans chaque espace interradial, on 
observe deux plaques successives principales, dont la plus externe est situee vers la 
peripherie du disque, avec d'autres plaques beaucoup plus petites. Les boucliers 
radiaux, de moyenne grosseur, sont triangulaires avec les angles arrondis, et un peu 
plus longs que larges : leur longueur ne depasse guere le quart du rayon du disque ; 
ils sont legerement divergents et separes sur toute leur longueur par la serie de plaques 
signalee plus haut. On trouve, sur leur bord libre, une rangee de papilles, petites, 
arrondies, peu developpees et formant une bordure reguliere dans les individus de 
taille moyenne ; dans les plus grands, ces papilles sont plus nombreuses, et elles sont 
disposees en deux ou meme en trois rangees d'ailleurs tres irregulieres : elles s'y 
montrent du reste moins developpees que dans les individus moyens. 

On remarque, sur les grands exemplaires, que les boucliers radiaux off rent, dans 
leur region centrale, une depression plus ou moins accentuee, tandis qu'ils se relevent 
vers leur bord externe en une ou deux preeminences arrondies. Cette disposition est 
analogue a celle que j'ai signalee chez VO. anceps, mais elle est moins accentuee. De 
plus, la plupart des plaques de la face dorsale du disque, surtout celles qui sont voisines 
des bords, ont une tendance a se soulever en une ou deux protuberances centrales : 
au moins sont-elles toujours plus ou moins convexes ; leur surface est rugueuse. 

La face ventrale du disque offre, vers la peripherie, une assez grosse plaque mediane 



ECHINODERMA 43 

arrondie, generalement plus large que longue, avec quelques autres plaques plus 
petites. Les plaques genitales sont allongees et elles portent, sur leur bord libre, une 
rangee de papilles courtes et peu developpees. Les fentes genitales sont etroites, mais 
elles s'etendent depuis 1'extremite des plaques adorales jusqu'a la peripherie du disque. 

Les boucliers buccaux sont assez grands, piriformes ou pentagonaux, relativement 
plus gros dans les individus de taille moyenne que dans les grands : dans ces derniers, 
ils sont piriformes avec le bord externe convexe ; dans les moyens, ils sont plutot 
pentagonaux et ofErent un angle proximal aigu limite par deux cotes droits et deux 
bords lateraux droits se reliant par des angles arrondis au cote distal convexe ; ils 
sont un peu plus longs que larges. Les plaques adorales sont allongees et etroites, 
avec les bords paralleles, beaucoup plus longues que larges. Les plaques orales sont 
egalement allongees et plus longues que larges, mais elles sont plus petites et un peu 
plus etroites que les precedentes. Les papilles buccales sont au nombre de quatre de 
chaque cote ; elles sont petites, basses, rectangulaires et subegales. La papille 
terminale impaire est petite et elle ne depasse guere la taille des precedentes. 

Les plaques brachiales dorsales sont assez grandes, quadrangulaires, avec un cote 
proximal etroit, un cote distal tres large et fortement convexe et des bords lateraux 
divergents et droits ; elles sont toutes contigues. Les premieres sont un peu plus larges 
que longues, puis elles deviennent aussi longues que larges et finalement plus longues que 
larges. Elles sont carenees, et, de plus, elles portent, un peu en arriere du bord distal, 
chacune un gros tubercule arrondi d'autant plus saillant et plus marque que les exem- 
plaires sont plus gros; ces tubercules rendent la carene des bras encore plus apparente. 

La premiere plaque brachiale ventrale est assez grande, triangulaire, avec Tangle 
proximal arrondi et le bord distal un peu convexe ; elle est aussi large que longue ou 
un peu plus large que longue. Les suivantes sont assez grandes, triangulaires, avec 
un angle proximal plutot aigu, limite par des cotes droits et un bord distal legerement 
convexe ; elles sont plus larges que longues. Les premieres sont contigues, mais 
elles se separent au dela du disque. 

Les plaques laterales, peu proeminentes, portent, a la base des bras, quatre petits 
piquants papilliformes ; ce chiffre tombe a trois a une certaine distance de la base 
des bras, et cela d'autant plus vite que 1'echantillon est plus petit. Les deux piquants 
ventraux sont tres rapproches 1'un de 1'autre et le suivant est un peu eloigne du precedent ; 
le dernier est tout a fait dorsal et place a une certaine distance du troisieme. 

Les pores tentaculaires sont peu developpes. Ceux de la premiere paire ne 
s'ouvrent pas dans la bouche : ils ofErent, sur leur bord externe, trois papilles basses, 
et, sur leur bord interne, deux ou trois papilles moins developpees que les precedentes, 
plus etroites et formant une bordure tres mince dans laquelle il est difficile de distinguer 
les limites des papilles. Les pores de la deuxieme paire sont tres petits et ils portent 
deux petites ecailles papilliformes ; on distingue encore les pores de la troisieme paire 
qui sont tres reduits et n'ont que deux ecailles rudimentaires, mais, au dela, les pores 
cessent d'exister. 



44 R. K(EHLER 

Rapports et differences. I/O. resistens rappelle beaucoup VO. anceps que j'ai 
decrite d'apres les exemplaires decouverts par la Scotia, par 71 lat. S. et 16 long. W.. 
a une profondeur de 1410 brasses, mais elle s'endistingue nettement par les fentes genitales 
plus allongees et par la presence de trois paires de pores tentaculaires : bien que les deux 
dernieres paires soient peu developpees, elles sont cependant bien visibles, tandis que 
YO. anceps n'en possede pas plus de deux paires en tout. Les plaques dorsales du 
disque sont moins epaisses dans YO. resistens que dans YO. anceps et si leur region 
centrale est deprimee, elles n'oflrent pas, comme dans cette derniere espece, cet epais- 
sissement peripherique que j'ai indique; enfin les piquants brachiaux sont au nombre 
de quatre a la base du bras. 

L'O. resistens s'eloigne de YO. martensi Studer, par les pores tentaculaires et par 
la forme des plaques brachiales ventrales. 



OPHIOGLYPHA JXEXIBILIS, nov. sp. (PI. V, fig. 1 et 2) 
Cap Royds. Profondeur, 60-80 brasses. 20 aout 1908. Quelques echautillons. 

Dans les plus grands individus, le diametre du disque egale 8 niillim. et les bras 
ont de 20 a 25 millim. de longueur ; dans les autres, qui sont les plus nombreux, le 
diametre du disque varie entre 3 et 6 millim. 

Le disque est arrondi ou subpentagonal ; la face dorsale est fortement convexe 
et la face ventrale est plane : il est epais, tandis que les bras sont au contraire tres 
greles, minces et flexibles. 

La face dorsale du disque est couverte de plaques tres inegales, a limites nettement 
indiquees. On distingue une grande centro-dorsale arrondie, et cinq radiales priiuaires 
egalement arrondies et de meme taille que la centro-dorsale ; les radiales sont separees 
les unes des autres par une rangee de petites plaques et le cercle qu'elles forment est 
aussi separe de la centro-dorsale par une rangee de plaques inegales, celles qui 
correspondent a 1'intervalle des radiales etant beaucoup plus grandes que les autres. 
En dehors de ces plaques, on reconnait dans les espaces radiaux une plaque triangulaire, 
plus grande que les voisines et qui separe les regions proximales des boucliers radiaux 
de chaque paire ; dans les espaces interradiaux, on remarque, vers la peripherie du 
disque. deux plaques tres grandes, arrondies, dont la derniere occupe le bord du disque. 
Les autres plaques sont beaucoup plus petites, inegales et irregulierement arrondies. 
Les boucliers radiaux sont petits et leur taille est inferieure a celle des grandes plaques 
du disque : ils sont triangulaires, divergents, separes sur toute leur longueur par deux 
plaques, la distale tres petite, la proximale grande et triangulaire ; ils sont raproches 
1'un de 1'autre au niveau de leur angle externe, mais non contigus sur les plus grands 
exemplaires. Ils sont a peine plus longs que larges et leur longueur ne depasse pas 
le quart du rayon du disque. En dehors de chaque bouclier, on observe une rangee 
de papilles assez basses, rectangulaires, obtuses ou legerement amincies a 1'extremite. 



ECHINODERMA 45 

La face ventrale n'offre, en dehors des grands boucliers buccaux, que quelques 
plaques peu nombreuses : on distingue une grande plaque mediane faisant suite 
au bouclier buccal et quelques autres plaques plus petites. Les plaques genitales 
sont etroites et elles portent sur leur bord libre une rangee de papilles basses. Les 
fentes genitales sont tres apparentes. 

Les boucliers buccaux sont grands, de forme pentagonale, aussi larges que longs : 
Tangle proximal, obtus, est limite par deux cotes jjpits et les bords lateraux se relient 
par deux angles arrondis au cote distal qui est convexe. Les plaques adorales sont 
assez petites, etroites, deux fois plus longues que larges. Les plaques orales sont 
petites et triangulaires. Les papilles buccales laterales ne depassent generalement 
pas le nombre de trois, elles sont rectangulaires, assez petites et elles font suite 
directement aux papilles internes du pore tentaculaire buccal ; la papille terminale 
impaire est un peu plus grande que les autres. 

Les plaques brachiales dorsales sont assez grandes. Les premieres sont rec- 
tangulaires et plus larges que longues, avec un cote proximal tres etroit, deux bords 
lateraux divergents et un cote distal tres large et fortement convexe qui se decompose 
ordinairement en trois cotes distincts se reunissant par des angles obtus. Le cote 
proximal ne tarde pas a disparaitre sur les plaques suivantes qui deviennent alors 
triangulaires en jneme temps qu'elles se montrent un peu plus longues que larges. 
Elles sont contigues sur le premier tiers des bras, puis elles se separent par un intervalle 
etroit et elles deviennent bientot un peu plus longues que larges. 

La premiere plaque brachiale ventrale est triangulaire, un peu plus large que 
longue, avec Tangle proximal legerement tronque ; le bord distal, tres convexe et 
offrant en son milieu un lobe assez large plus ou moins apparent, se decompose parfois 
en trois petits cotes distincts. La deuxieme plaque est rectangulaire, un peu plus 
longue que large, avec un cote proximal plus etroit, un cote distal large et convexe et 
des bords lateraux divergents. Les suivantes sont pentagonales avec un angle proximal 
aigu, et un bord distal large et convexe offrant en son milieu un petit lobe plus ou 
moins accuse ; ces plaques se separent a partir de la troisieme. 

Les plaques laterales, peu proeminentes, portent chacune quatre piquants courts, 
assez larges, coniques et pointus, s'inserant le long du bord distal de la plaque a des 
interval les egaux. 

Les pores tentaculaires ne sont pas tres developpes. Ceux de la premiere paire, 
qui s'ouvrent dans la bouche, portent sur chacun de leurs bords trois ecailles rec- 
tangulaires et obtuses, les internes plus fortes que les externes. Les pores de la 
deuxieme paire portent trois ecailles proximales et externes, epaisses, coniques, pointues, 
et deux ecailles distales et internes tres petites, basses et peu developpees ; ceux de 
la troisieme paire out trois ecailles proximales et une ou deux distales ; ceux de la 
quatrieme paire out deux ou trois ecailles proximales et une seule distalc- ; enfin les 
pores de la cinquieme paire ont deux ecailles proximales et une distale. Au dela, il 
n'existe qu'une seule ecaille proximale. 



46 R. KCEHLER 

Dans les jeunes exemplaires, le nombre des plaques dorsales du disque est beaucoup 
moins eleve : les six plaques primaires, avec les deux grandes plaques de chaque inter- 
radius, suffisent pour recouvrir la plus grande partie de la face dorsale du disque. Les 
boucliers radiaux sont rapproches en dehors et parfois meme contigus sur une certaine 
partie de leur longueur. 

Rapports et differences. L'O. flexibilis se rapproche de VO. mimana que j'ai decrite 
d'apres des exemplaires recueillis par la Scotia, a une profondeur de 1410 brasses, par 
71 22' lat. S. et 16 34' long. W. ; elle en differe par ses plaques brachiales ventrales 
qui deviennent rapidement plus longues que larges, tandis que chez YO. mimaria elles 
restent toujours beaucoup plus larges que longues, par la face ventrale du disque 
offrant une grande plaque mediane faisant suite au bouclier buccal et en dehors de 
laquelle il n'existe qu'un petit nombre de plaques seulement, enfin par les piquants 
brachiaux qui ne sont qu'au nombre de quatre. 

Elle s'ecarte de I'O. martensi Studer, de la Georgie du Sud, par une forme complete- 
ment differente des plaques brachiales dorsales et ventrales, ainsi que par les pores 
tentaculaires, et par la presence de quatre piquants brachiaux. 

AMPHIURA ALGIDA, nov. sp. (PI. VII, fig. 14 et 15) 

Cap Koyds. Profondeur, 10-20 brasses. Deux echantillons. 
Cap Royds. Profondeur, 50-80 brasses. Un echantillon. 

Dans le plus grand individu, le diametre du disque est de 4*5 millim. ; dans les 
deux autres, il ne depasse pas 3 millim. Les bras sont tous casses et ils ne devaient pas 
avoir plus de 15 millim. de longueur. 

Le disque est pentagonal avec les angles arrondis. La face dorsale est couverte 
de plaques grandes et inegales, en forme d'ecailles aplaties, imbriquees, dont le bord 
libre offre une tres mince bordure, sorte de lisere finement strie et transparent. Ces 
plaques sont tres grandes dans la region centrale du disque : elles deviennent plus 
petites vers la peripherie et vers les boucliers radiaux ; il n'y a pas la moindre indication 
de plaques primaires. Les boucliers radiaux sont petits et peu developpes : ils sont 
a peine deux fois et demie plus longs que larges et leur longueur est inferieure au quart 
du rayon du disque ; ils sont assez rapproches 1'un de 1'autre, peu divergents et 
separes par une rangee etroite de plaques. 

La face ventrale du disque est recouverte de plaques identiques a celles de la face 
dorsale, mais plus petites et egales. Les fentes genitales sont etroites. 

Les boucliers buccaux sont triangulaires avec des cotes convexes et un angle proximal 
arroudi ; le cote distal est fortement convexe et il offre meme en son milieu un lobe 
qui n'estpas tres preeminent dans le grand exemplaire ou le bouclier est a peupres aussi 
long que large ; dans les deux petits, ce lobe est au contraire tres preeminent et les 
boucliers sont plus longs que larges. Les plaques adorales sont triangulaires, avec 



ECHINODERMA 47 

le bord oral legerement incurve ; elles sont fortement amincies en dedans et a peine 
contigues sur la ligne mediane : parfois meme elles restent separees de leurs congeneres 
par Tangle proximal du bouclier buccal ; leur region externe est au contraire fortement 
developpee et arrive meme a toucher la region correspondante de la plaque opposee 
en comprimant la premiere plaque brachiale ventrale. Les plaques orales sont 
etroites et assez hautes. Les papilles buccales comprennent d'abord deux papilles 
placees sur le meme plan que les autres pieces buccales : 1'interne est epaisse et arrondie, 
1'externe est allongee, proeminente, assez epaisse et son extremite est obtuse ; il existe 
en outre, sur un plan superieur, une petite papille conique et pointue s'inserant entre 
les deux precedentes. 

Les plaques brachiales dorsales sont grandes et triangulaires avec les cotes legere- 
ment convexes et un angle proximal arrondi ; elles sont un pen plus larges que longues 
et toutes contigues. 

La premiere plaque brachiale ventrale est tres petite et triangulaire, avec un angle 
distal aigu qui est limite de chaque cote par les plaques adorales. Quand ces dernieres 
plaques sont tres rapprochees ou contigues, Tangle de la plaque, resserre entre elles, 
est tres pointu ; quand les plaques adorales sont un peu ecartees, Tangle devient tronque. 
Les plaques suivantes sont pentagonales, avec un angle proximal : sur les premieres 
plaques, qui sont a peu pres aussi longues que larges, cet angle est d'abord tres obtus ; 
il se montre plus aigu sur les plaques suivantes qui deviennent sensiblement plus 
longues que larges. Le bord distal s'echancre legerement au-dela du disque. Toutes 
ces plaques sont contigues. 

Les plaques laterales portent quatre piquants subegaux et dont la longueur egale 
celle de Tarticle ; ces piquants, assez epais, s'amincissent progressivement jusqu'a 
Textremite qui forme une pointe obtuse. 

L'ecaille tentaculaire, unique, est bien developpee ; elle offre deux cotes paralleles et une 
extremite obtuse : elle est presque rectangulaire et pres de deux fois aussi longue que large. 

Rapports et differences. 1>'A. algida se distingue facilement de toutes les 
Amphiura s. str. connues possedant des ecailles sur les deux faces du disque et une 
seule ecaille tentaculaire. L'.4. prcefecta, que j'ai decrite d'apres un exemplaire de 
Tile Campbell, a Tecaille tentaculaire remarquablement grande et plus forte que chez 
\'A. algida ; les plaques primaires sont tres apparentes, la papille buccale externe est 
squamiforme et la face ventrale du disque est couverte d'ecailles tres petites. 
L ; 'A. pusilla Farquhar, de la Nouvelle-Zelande, a six piquants brachiaux et son ecaille 
tentaculaire, arrondie, est assez petite. ISA. magellanica, avec son premier piquant 
ventral allonge, est bien differente de VA. algida. 

Notre espece se rapproche surtout de \'A. angularis Lyman, mais, chez cette 
derniere, la face ventrale du disque est nue ou garnie d'ecailles rudimentaires ; de 
plus, la papille buccale externe est conique et pointue, Tecaille tentaculaire est arrondie 
et les boucliers buccaux sont presque circulaires. 



48 R. KCKHLER 

OPHIODIPLAX, nov. gen. 

Ce genre rappelle les OphiacantJm et les genres voisins. 

La face dorsale du disque est recouverte d'un tegument qui cache completement les 
plaques sous-jacentes, y compris les boucliers radiaux, et qui porte de petits batonnets ; 
ce tegument se continue sur les premieres plaques brachiales dorsales. Celles-ci sont 
divisees, par une suture transversale, en deux moities inegales, la proximale plus 
petite, et cette division existe sur toute la longueur des bras. Les papilles buccales 
sont nombreuses et elles ne forment pas une rangee reguliere. 

La division en deux des plaques brachiales dorsales s'observe d'une maniere tres 
constante dans les nombreux exemplaires que j'ai pu etudier, aussi bien chez les 
individus tres jeunes que chez ceux dont le disque atteintlO a 11 millim. de diametre 
et qu'on peut considerer comme adultes. Ce caractere ne s'observe chez aucune 
Ophiacanthidee connue et il m'a paru suffisant pour justifier la creation d'un nouveau 
genre. 

OPHIODIPLAX MSJUNCTA, nov. sp. (PL VI, figs. 9, 10 et 11 ; PI. VII, fig. 13) 
Cap Royds. Profondeur, 60-80 brasses. 20 aout 1908. Trois echantillons. 

Le diametre du disque varie entre 8 et 9 millim. ; les bras atteignent 40 a 50 millim. 
de longueur. 

Le disque est subpentagonal dans 1'un des individus ; dans les deux autres, il est 
pentagonal avec les cotes plus ou moins excaves. 

La face dorsale du disque est couverte d'un tegument qui cache completement les 
plaques sous-jacentes et offre de petits batonnets assez ecartes, courts, coniques et 
dont le sommet, emousse, porte de deux a quatre ou cinq spinules tres fines et pointues. 
A la peripherie du disque, ces batonnets s'allongent un peu et deviennent cylindriques ; 
leur surface est plus ou moins rugueuse et ils montrent toujours un certain nombre de 
petites spinules a leur partie terminale. Les boucliers radiaux ne sont pas distincts : ils 
sont settlement indiques par des saillies plus ou moins apparentes de la face dorsale du 
disque a la base des bras. 

La face ventrale du disque presente des batonnets identiques a ceux de la face 
dorsale, mais plus epais que sur cette derniere, surtout au voisinage des fentes genitales 
ou Ton distingue des plaques tres fines et arrondies ; ces batomiets se continuent, en 
diminuant de longueur, jusqu'au voisinage des boucliers buccaux. Les fentes genitales 
sont tres larges. 

Les boucliers buccaux sont assez grands, quadrangulaires, plus larges que longs, 
avec les angles arrondis ; la region proximale, qui correspond au sommet de Tangle 
proximal, est parfois separee du reste de la plaque par un sillon irregulier. Les plaques 
adorales sont assez larges mais pas tres grandes, et elles sont a peine deux fois et demie 
plus longues que larges ; elles ne possedent pas de lobe distal separant le bouclier 



ECHINODERMA 49 

buccal de la premiere plaque brachiale ventrale. Les plaques orales sont petites et 
triangulaires. Les papilles buccales sont nombreuses et serrees, et elles ne forment 
pas une serie tres reguliere, car, dans la region moyenne, on observe deux rangees 
plus ou moins distinctes ; il y en a une.dizaine en tout de chaque cote. Ces papilles 
sont petites, coniques et pointues, et elles deviennent un peu plus longues vers 
Fextremite proximale des plaques orales ; la papille impaire terminale est un peu 
plus grande et plus forte que les voisines. 

Les contours des premieres plaques brachiales dorsales ne sont pas distincts. Le 
tegument de la face dorsale du disque se continue, en efiet, sur une certaine longueur 
de la face dorsale des bras en recouvrant les cinq ou six premieres plaques brachiales : 
il presente des batonnets analogues a ceux de la face dorsale du disque, mais ceux-ci 
sont plus courts, coniques avec la pointe rugueuse ou garnie de deux ou trois petites 
spinules, et ils deviennent de plus en plus petits pour disparaitre finalement. Les 
plaques brachiales dorsales apparaissent alors et elles se montrent de suite avec leur 
structure caracteristique : chacune d'elles est en effet divisee par un sillon transversal 
legerement convexe en deux parties inegales ; la partie proximale est petite et la partie 
distale est relativement grande. Dans leur ensemble, ces plaques ont une forme 
triangulaire et elles sont un peu plus longues que larges avec un sommet proximal 
arrondi et un bord distal convexe. La region proximale de la plaque est petite, en 
forme de triangle a sommet plus ou moins arrondi : cette region est un peu plus longue 
que large. La region distale de la plaque est plus large que longue, trapezoiidale, 
avec un cote proximal etroit et un peu concave, un cote distal tres large et convexe 
et deux bords lateraux divergents. 

La premiere plaque brachiale ventrale est assez grande, triangulaire, avec les 
angles arrondis ; le sommet, distal, est arrondi et la base, proximale, est convexe ; 
elle est un peu plus large que longue. Les plaques suivantes sont triangulaires, plus 
larges que longues, avec un angle proximal obtus et arrondi, et un bord distal forte- 
ment convexe ; ce bord peut offrir un petit lobe median obtus et il se decompose 
parfois en deux cotes distincts. Ces plaques sont separees des la base du bras. A 
partir du premier tiers, elles deviennent plus longues que larges, avec un angle 
proximal aigu un peu arrondi et un bord distal toujours fortement convexe. 

Les plaques laterales, assez proemineutes, portent six piquants a la base des bras : 
la longueur de ces derniers augmente legerement depuis le premier, qui egale Farticle, 
jusqu'au troisieme ; le quatrieme et le cinquieme sont un peu plus longs et leur 
longueur depasse un article et demi ; le sixieme est un peu plus court. Leur surface 
est couverte de tres fines rugosites qui sont un peu plus marquees vers 1'extremite. 
On trouve parfois sept piquants a une petite distance de la base des bras. 

L'ecaille tentaculaire, unique, est courte, mince, spiniforme et son extremite est 
pointue. 

J'ai rencontre dans les collections de la deuxieme Expedition Antarctique du 

BK1T. ANTARCT. EXPED. 1907-9. VOL. II. H 



50 R. KXEHLER 

Dr. Charcot, un certain nombre d'exemplaires de cette meme Ophiure qui ni'ont 
permis de mieux en preciser les caracteres et de m'assurer de leur Constance ; j'ai 
note egalement quelques particularites qu'il est bon de signaler. La taille des 
echantillons peut etre superieure a celle de ceux qui ont ete recueillis par 1'Expedition 
Anglaise, et, dans certains d'entre eux, le diametre du disque depasse 11 millim. J'ai 
represente ici deux de ces individus ; dans 1'un, qui est vu par la face dorsale, le 
diametre du disque atteint a peine 9 millim. (PI. VI, fig. 10) ; dans Fautre, qui est 
vu par la face ventrale, le diametre du disque est de 1T2 millim. (PI. VI, fig. 11). Les 
plaques brachiales dorsales sont toujours divisees, mais leur forme peut offrir des 
variations : tantot elles sont plus longues que larges comme dans 1'individu reproduit 
PI. VI, fig. 10, tantot au contraire elles sont un peu plus larges que longues. 
Les piq uants brachiaux peuvent etre plus developpes et le piquant dorsal depasse 
la longueur de deux articles, ainsi que cela arrive dans 1'exemplaire de la fig. 10, 
dont le disque n'a que 9 millim. Dans les echantillons de grande taille (PI. VI, 
fig. 11), les papilles buccales deviennent plus fortes et plus nombreuses ; les plaques 
brachiales ventrales offrent, a partir de la deuxieme, une forme pentagonale avec un 
angle proximal tres obtus et un bord distal convexe ; ce n'est qu'a partir de la dixieme 
qu'elles deviennent triangulaires. D'une maniere generale, je remarque que, sur les 
exemplaires de I'Expedition Charcot, les piquants de la face dorsale du disque se 
continuent moins loin sur les bras que sur les individus de la Terre Victoria et ils 
peuvent disparaitre des la troisieme plaque brachiale dorsale. 

L'O. disjuncta me parait identique a une Ophiure antarctique que J. Bell a 
figuree, sans la decrire (08, PL IV) : si elle n'est pas identique a cette forme, elle en est, 
en tout cas, fort voisine. D'apres les dessins de J. Bell, les plaques brachiales dorsales 
de cette Ophiure sont divisees, a la base des bras, en deux parties inegales ; mais, 
d'autre part, les pores tentaculaires auraient chacun deux ecailles, 1'une proximale, 
1'autre distale, et ces ecailles sont elargies transversalement ; je n' observe aucune 
disposition analogue sur mes echantillons et je me demande s'il n'y a pas une erreur 
dans les dessins de J. Bell. J'ajouterai que cet auteur considere son Ophiure comme 
un jeune : or sur deux dessins qui sont faits avec un grossissement de 9/4, le diametre 
du disque est de 22 millim. mesure entre le fond d'un espace interradial et le bord 
radial oppose, ce qui correspond a un diametre de pres de 10 millim. en grandeur 
naturelle. Si done ces chiffres sont exacts, les exemplaires de Bell seraient de la taille de 
ceux que j'ai etudies et il me parait bien difficile d'admettre qu'une Ophiacanthidee, 
dont le disque a un diametre moyen de 10 millim., soit un jeune. 

Le savant naturaliste anglais parait attribuer a cette jeunesse des exemplaires les 
caracteres des plaques brachiales dorsales qui ne seraient pas completement calcifiees. 
Je ne suis pas de cet avis : j'estime que ces plaques sont parfaitement formees et 
qu'elles restent fragrnentees chez 1'adulte, comme on 1'observe d'ailleurs chez quelques 
autres Ophiures. 



ECHINODERMA 51 

ECHINIDES 

STERECHINUS NEUMAYERI (Meissner) 
Voir pour la bibliographie : Mortensen (10), p. 64. 

Cap Royds. Profondeur, 10-50 brasses. Juillct ^1908. Quelques echantillons de 
differentes dimensions : le diametre du test varie entre 58 et 24 millim. 

ABATUS SHACKLETONI, nov. sp. (PL IV, fig. 3 a 10 ; PL VIII, fig. 1 a 6) 

Baie du Cap Royds. 5 ma-rs 1908. Profondeur, 10-20 brasses. Treize 
echantillons dont un incomplet et casse. 

Cinq exemplaires sont d'assez grande taille et leur longueur est voisine de 35 millim., 
les autres sont plus petits. Dans les plus grands, la longueur totale avec les piquants 
varie de 34 a 38 millim. 

Dans 1'individu represents PL VIII, fig. 3 a 5, les dimensions sont les suivantes : 

Longueur (sans les piquants) 36 millim. 

Largeur ..... 31 

Hauteur ,, . .... 19*5 

Get individu a les poches incubatrices tres profondes et il represents incontestablement 
une femelle. On voit, par les photographies que j'en donne, que sa forme est reguliere- 
ment ovoiide et que le corps est assez allonge. Cette meme forme s'observe sur trois 
des autres grands echantillons et me parait etre typique. Le dernier exemplaire est 
relativement plus large et plus court, ainsi qu'on peut le constater sur la photo- 
graphic reproduite PL VIII, fig. 6. Les dimensions de cet individu sont les suivantes : 

Longueur (sans les piquants) ..... 33 millim. 
Largeur ,, ..... 32 

Hauteur ,, ..... 19 ,, 

Les poches incubatrices de cet exemplaire sont beaucoup moins profondes que celles 
du precedent, mais je le considere neanmoins comme une femelle. Un trois'eme 
individu, dont j'ai laisse le test intact, presente des poches incubatrices profondes et 
appartient aussi au sexe femelle. Quant aux deux autres individus, ils ont les 
ambulacres dorsaux a peine deprimes et non transformes en poches incubatrices : ce 
sont des males. 

Les autres exemplaires, de plus petite taille, ont le corps relativement elargi et 
moins allonge ; leurs ambulacres dorsaux sont peu deprimes et tous ont le caractere 
de males. 



52 R. K(EHLER 

Voici les dimensions que je releve, piquants non compris, sur les sept echantillons 
qui sont intacts : 

L&n^ueur. Largeur. 

13 millim. 11' 5 millim. 

16 13 

17 14 
17 15-5 
21-5 19 

22 19-5 

23 19 

Je decrirai 1'exemplaire represente PL VIII, fig. 3 a 5, que je prendrai comme type de 
1'espece ; j'indiquerai ensuite les quelques differences que je releve avec celui de la 
PI. VIII, fig. 6, et enfin je donnerai les caracteres du male. 

Vu d'en haut, le test est assez regulierement ovoide ; il est plus retreci en arriere 
qu'en avant et la plus grande largeur s'observe au niveau de Fappareil apical qui est 
plus rapproche du bord anterieur que du bord posterieur. 

Le bord anterieur est a peine deprime sur la ligne mediane, le sillon qui correspond 
a 1'ambulacre dorsal anterieur devenant de moins en moins profond a mesure qu'on 
se rapproche de 1'ambitus ; nieme 1'echancrure, qui, dans le premier exemplaire, est 
a peine marquee en dessous du fasciole, entre celui-ci et 1'ambitus, devient, ainsi que 
nous le verrons plus loin, tout a fait nulle dans le second ou le sillon est d'ailleurs moins 
profond, comme cela arrive aussi chez les males. 

Vu lateralement, le test offre d'abord une courbe s'elevant regulierement jusqu'a 
1'appareil apical, puis, en arriere de cet appareil, il forme une preeminence interradiale 
assez marquee sur I'echantillon depourvu de ses piquants, mais qui n'apparait pas ou 
n'apparait que fort peu sur les individus munis de leurs piquants ; cette preeminence 
diminue rapidement jusqu'a 1'extremite posterieure qui est tronquee verticalement et 
peu elevee. La face ventrale est tout a fait plane en avant du peristome et un peu 
convexe en arriere. 

L'appareil apical est situe en avant du milieu du corps et il presente trois orifices 
genitaux, deux a gauche et un a droite ; ces orifices sont arrondis et assez grands 
(PI. VII, fig. 4). Les deux orifices de gauche sont tres rapproches 1'un de 1'autre 
tandis que 1'orifice de droite est assez ecarte de 1'orifice posterieur gauche : il n'est 
pas situe exactement sur le meme niveau transversal que ce dernier, mais se trouve 
place tres legerement en arriere. La ligne qui reunit les deux orifices passe a 15' 5 millim. 
du bord anterieur du test eta 20' 5 millim. du bord posterieur. La plaque madreporique 
occupe a peu pres tout 1'espace compris entre les deux orifices posterieurs ; elle s'etend 
en avant jusqu'au niveau du bord anterieur de 1'orifice anterieur, et, en arriere, elle 
depasse le bord posterieur des orifices posterieurs. Elle est criblee de petits pores 
arrondis, regulierement disposes et qui en occupent a peu pres toute la surface ; on 



ECHINODERMA 53 

reconnait, en outre, un certain nombre de granules tres fins et places aussi d'une 
maniere reguliere. On distingue facilement les contours des plaques ocellaires qui 
sont petites, pentagonales avec un angle proximal et offrcnt chacune un orifice trans- 
versal en forme de croissant a concavite tournee vers 1'exterieur. Les contours des 
autres plaques sont indistincts. 

L'ambulacre anterieur impair forme, a la face dorsale du test, un sillon assez etroit 
et peu profond, dont les cotes se continue nt paimn bord tres arrondi avec les regions 
interradiales du test et qui s'attenue progressivement avant d'atteindre le fasciole ; 
il disparait presque complete ment au niveau du bord anterieur du test (PL VIII, fig. 5). 
Les zones poriferes, situees sur les cotes legerement obliques du sillon, sont droites 
et elles vont en divergeant quelque peu ; elles ne sont pas tres ecartees 1'une de 
1'autre en raison de la faible largeur du sillon. Je compte dans chacune d'elles, et 
jusqu'au fasciole, vingt paires de pores dont la grosseur augmente depuis la premiere 
jusqu'a la septieme, puis decroit ensuite de telle sorte que les pores des deux dernieres 
paires sont tres fins. Les huit ou dix premieres paires de pores sont separees par deux 
ou trois granules formant souvent une petite rangee trans versale ; au dela, les 
granules diminuent ou meme disparaissent completement entre les pores, tandis qu'on 
voit apparaitre sur chaque plaque un granule plus gros et rapproche de son bord 
interne. Les pores des premieres paires sont disposes suivant une ligne perpendicu- 
laire a 1'axe du sillon, puis les paires suivantes se placent obliquement par rapport a 
cet axe, en meme temps que les pores de chaque paire se rapprochent 1'un de 1'autre 
et deviennent plus petits. 

Les ambulacres lateraux anterieurs et posterieurs presentent a peu pres le meme 
developpement. Les ambulacres anterieurs sont fortement divergents et ils sont 
presque places sur le prolongement 1'un de 1'autre, tandis que les posterieurs sont 
tres rapproches et peu divergents ; ils sont separes par la preeminence interradiale 
que j'ai signalee plus haut et dont la largeur, a la base, est de 4 millim. L'axe de 
1'ambulacre anterieur forme, avec celui de 1'ambulacre posterieur du meme cote un 
angle de 90 environ. 

Les ambulacres anterieurs sont, comme les posterieurs, tres profondement 
deprimes a la face dorsale du test et ils constituent des poches incubatrices, allongees 
mais assez etroites dans notre espece ; elles ont toutes les memes dimensions et 
mesurent environ 12 millim. de longueur sur 3 millim. de largeur ; les poches 
anterieures sont legerement elargies sur leur region externe, tandis que les poches 
posterieures conssrvent la meme largeur sur toute leur etendue. La depression qui 
constitue chaque poche se fait tres brusquement a partir de la troisieme paire de pores 
ambulacraires et les parois sont a peu pres verticales ; elles se relient par un angle 
arrondi avec la face dorsale du test. La profondeur des poches est de 6 millim. La 
partie de 1'ambulacre qui est comprise entre 1'appareil apical et la poche proprement 
dite, ne reste pas a fleur du test, mais elle est un peu deprimee surtout dans le sens de 
la longueur de maniere a former un sillon oblique qui conduit a la poche. Chaque 



54 R. KCEHLER 

poche s'etend jusqu'au fascicle : en 1'abordant, elle se releve brusquement pour la 
poche posterieure et un peu obliqucment pour 1'anterieure. 

La region interradiale dorsale comprise entre le sillon et 1'ambulacre lateral 
anterieur est convexe ; elle se continue insensiblement avec le sillon tandis qu'elle 
s'arrete brusquement au bord de la poche incubatrice. Elle est recouverte de gramiles 
primaires de petite taille, de dimensions tres uniformes et assez espaces ; ces granules 
sont plus serres au voisinage de la poche incubatrice et leurs dimensions augmentent 
un peu au bord meme de cette poche. Entre ces granules primaires se montrent 
d'autres granules extremement fins. On retrouve ces deux sortes de granules sur les 
regions interradiales anterieures et posterieures, ainsi que sur la preeminence inter- 
radiale posterieure, et Ton constate que les granules primaires deviennent toujours 
un peu plus serres et un peu plus gros au voisinage des poches incubatrices. 

Le face ventrale est peu convexe (PI. VIII, fig. 3). L'arnbulacre anterieur est peu 
distinct: il est legerement deprime au voisinage du peristome, mais il se releve en 
meme temps qu'il se retrecit a mesure qu'on se rapproche du bord du test. II offre, 
de chaque cote, quatre paires de pores ambulacraires rapproches, entoures chacun 
d'un cercle ovalaire tres apparent ; ca et la se montre un granule extremement fin. 
Puis les paires s'espacent rapidement en meme temps que les pores deviennent plus 
fins : les granules au contraire deviennent subitement plus gros et leurs dimensions 
depassent celles des granules dorsaux auxquels ils passent d'ailleurs a 1'ambitus. Les 
ambulacres lateraux anterieurs sont places sur le prolongement Fun de 1'autre de part 
et d'autre de la bouche ; ils offrent, de chaque cote, cinq paires de pores, entoures 
chacun de leur cercle ovalaire, formant deux rangees convergentes separees par quelques 
fins granules ; puis les pores deviennent tres fins et espaces, tandis que les granules 
acquierent les memes dimensions que dans les interradius voisins. Les ambulacres 
lateraux posterieurs constituent de larges avenues n'offrant que des granules tres fins 
et peu abondants ; des granules plus gros et plus serres n'apparaissent qu'au voisinage 
de 1'extremite posterieure. 

Les regions interradiales anterieures et posterieures de la face ventrale sont 
uniformement couvertes de granules assez espaces, de taille uniforme, mais 
sensiblement plus gros que ceux de la face dorsale avec lesquels ils se continuent 
a 1'ambitus ; entre ces granules primaires on en voit d'autres tres fins et peu 
serres. Le plastron sternal est simplement convexe et il est arrondi a son extremite 
posterieure, sans former de preeminence ou de pointe saillante. II offre des granules 
disposes en rangees obliques divergentes partant de son extremite posterieure et se 
dirigeant veis les avenues ambulacraires ventrales : ces granules sont d'abord tres 
fins, mais leurs dimensions augmentent a mesure qu'on se rapproche des avenues 
ambulacraires et du peristome. 

Le peristome se trouve a peu pres a egale distance entre le bord anterieur du test 
et le milieu de la face ventrale; il est assez profondement situe, etroit, en forme de 
croissant, et plutot petit. Le labre est de dimensions moyennes ; son bord anterieur 



ECIIINODERMA 55 

est fortement convexe et il forme une saillie tres marquee, en forme de bee preeminent, 
qui cache la plus grande partie du peristome ; ses bords lateraux sont excaves et le 
bord posterieur, fortement arrondi et convexe, s'etend jusqu'au milieu de la deuxieme 
plaque ambulacraire voisine. 

La face posterieure est verticale, mais peu etendue ; elle est en grande partie 
occupee par le periprocte (PL IV, fig. 6). Celui-ci, de taille moyenne, a une forme 
ovo'ide un peu irreguliere ; il est un peu plus haiit_que large et mesure 5'1 millim. sur 
4'6 ; il ofire une rangee peripherique de grandes plaques rectangulaires et deux cercles 
plus ou moins reguliers de plaques centrales plus petites. II n'y a pas la moindre 
indication de tubes ambulacraires au voisinage du periprocte. 

Le fascicle offre un contour assez regulier, sans inflexions ni sinuosites bien 
marquees (PI. IV, fig. 6; PI. VIII, fig. 4 et 5). II est tres rapproche de 1'ambitus 
entre 1'ambulacre anterieur et la poche incubatrice anterieure a I'extremite de laquelle 
il est tangent : dans ce parcours il a la forme d'un arc de cercle presque regulier, a part 
deux sinuosites tres legeres. Entre les deux poches incubatrices de chaque cote, le 
fascicle s'eearte progressivement de 1'ambitus et devient legerement concave ; au 
niveau de l'extremite de la poche posterieure, a laquelle il est egalement tangent, il 
presente un angle obtus tres marque et se dirige vers 1'interradius posterieur, en 
suivant une direction a peu pres parallele au bord posterieur du test : il reste separe 
de ce bord par un intervalle de 4 millim. environ. Le fasciole conserve la meme 
largeur sur tout son trajet, soit T5 millim. environ. 

Les tubercules primaires sont performs et finement creneles. Us sont plus petits 
sur la face dorsale et plus gros sur la face ventrale. Leurs dimensions restent 
uniformes sur presque toute 1'etendue de la face doisale, sauf au voisinage immediat 
des poches incubatrices ou ils deviennent un peu plus gros. Sur la face ventrale, ils 
sont plus espaces et plus gros, mais leur taille diminue a mesure qu'on s'eloigne du 
peristome et ils passent progressivement aux granules de la face dorsale. Au milieu 
de ces granules primaires s'en montrent d'autres tres fins, dont les dimensions restent 
uniformes sur les deux faces du test. 

Les piquants de la face dorsale sont fins et assez courts, leur longueur ne depassant 
pas 3 millim. ; ils sont couches sur le test en formant un feutrage serre ; ils sont 
cylindriques, obtus a l'extremite, et, vers 1'ambitus, ils sont legerement recourbes. 
Les piquants des bords des poches incubatrices sont plus longs, un peu aplatis et ils 
sont diriges de maniere a recouvrir ces poches qui sont a peine visibles sur les 
echantillons non depouilles de leurs piquants. Les piquants de la face ventrale 
sont bien developpes et leur longueur peut atteindre 5 millim. 

Les pedicellaires appartiennent aux trois types tridactyle, rostre et globifere. Les 
pedicellaires tridactyles sont de deux formes differentes, mais toujours a trois valves. 
Les uns rappellent ceux que Mortensen a decrits et figures chez YA. cavernosus (10 bis, 
PI. XIX, fig. 37) : les valves sont elargies en forme de cuillerons qui sont peu 
retrecis a leur base et qui se toucheut sur une bonne partie de leur longueur ; ces bords 



56 R. K(EHLER 

sont munis de denticulations extremement fines et regulieres. Les pedicellaires de 
la deuxieme forme ont les valves plus etroites, recourbees, separees sur la plus grande 
partie de leur longueur et contigues seulement dans leur partie terminale qui est 
elargie et munie de denticulations aigue's : cette forme se rapproche ainsi des pedi- 
cellaires rostres. Ces deux sortes de pedicellaires sont repandus sur tout le test, la 
premiere plus abondamment que la seconde ; toutes deux atteignent une assez grande 
taille et la longueur de la tete peut etre de 0'5 a 0'6 millim. 

Les pedicellaires rostres ressemblent a ceux que Mortensen a indiques chez 
I' A. cavernosus (10 bis, PL XIX, fig. 30, 38 et 45) ; ils ont des valves fortement 
recourbees, qui, a partir de la base triangulaire, restent etroites en s'amincissant fort 
peu jusqu'a leur extremite qui est obtuse. Les bords sont depourvus de denticula- 
tions mais ils sont un peu sinueux et ils se relevent, de distance en distance, en une 
dent peu saillante ; 1'extremite de la valve n'est pas denticulee. Ces pedicellaires 
sont plus rares que les precedents et ils restent generalement plus petits ; la longueur 
de leur tete peut cependant atteindre 0'3 a 0'4 millim. 

Les pedicellaires globiferes sont surtout repandus au voisinage du periprocte ; ils 
rappellent beaucoup ceux que j'ai decrits et figures chez YA. elongatus ( 08, p. 620, 
PI. XVI, fig. 154). Les valves sont plus recourbees que dans cette derniere espece ; 
elles se terminent, comme chez elle, par deux crochets extremement developpes, en 
arriere desquels se trouve une fente allongee a la suite de laquelle vient une courte 
partie tubulaire. Ces valves ressemblent aussi a celles des pedicellaires globiferes 
decrits par Mortensen dans la var. bidens de YA. cavernosus, mais, ici, la fente qui 
precede les deux crochets terminaux est plus allongee. Le tige calcaire offre un peu 
avant son extremite un tres leger epaississement parfois a peine marque. Les valves 
et I'extremite distale du pedoncule sont enveloppees, comme chez YA. elongatus, d'un 
tissu fortement pigmente et glandulaire qui rend la tete du pedicellaire fort apparente 
et dont 1'ensemble mesure un millimetre de longueur environ, tandis que les valves 
incluses dans cette enveloppe ne depassent pas 0'35 millim. 

Les spicules des tubes ambulacraires sont petits et de forme tres irreguliere. Les 
plus simples se presentent sous forme de batonnets plus ou moins recourbes et assez 
epais, offrant un nombre variable de prolongements lateraux en forme de dents ; 
quand ces prolongements se rejoignent, il en resulte la formation de petites plaques 
perforees, allongees et de forme tres irreguliere. 

Les gros exemplaires ont une couleur brunatre assez foncee et uniforme ; les 
petits sont d'un brun plus clair et certains d'entre eux sont simple ment grisatres. 

L'exemplaire qui est represente PI. VIII, fig. 6, difiere par quelques caracteres de 
celui que je viens de decrire. Ainsi que je 1'ai dit plus haut, le test est relativement 
elargi au niveau de 1'appareil apical et celui-ci est plus rapproche du bord anterieur : 
la ligne qui joint les centres des deux orifices genitaux posterieurs se trouve a 13 millim. 
du bord anterieur et 20 millim. du bord posterieur du test. Les orifices genitaux sont 
un peu plus petits que dans le type. Le sillon auterieur est moins profond et le bord 



ECHINODERMA 57 

anterieur du test cesse d'etre deprime en son milieu ; les pores ambulacraires du sillon 
sont plus fins. La preeminence interradiale dorsale, qui separe les deux ambulacres 
posterieurs, est aussi moins accusee ; enfin le periprocte est un peu plus petit : il est 
sensiblement plus haut que large et il mesure 5 millim. de hauteur sur 4'5 de largeur ; 
il se rapproche un peu de la forme que nous observerons chez le male. Les tubercules 
de la face dorsale du test sont repartis plus uniformement que dans le type et la 
difference de taille que Ton observe avec ceux qui avoisinent les ambulacres lateraux 
est moins apparente. 

Les ambulacres lateraux anterieurs et posterieurs de la face dorsale sont assez 
f ortement deprimes, mais les fossettes qu'ils constituent sont beaucoup moins profondes 
que dans le type et leur profondeur ne depasse pas 3'5 millim. Au lieu d'apparaitre 
brusquement et d'offrir un bord proximal abrupt et vertical, les fossettes se creusent 
progressivement par I'enfoncement graduel des ambulacres. Je compte dix-neuf paires 
de pores dans 1'ambulacre anterieur, depuis 1'extremite proximale de 1'ambulacre 
jusqu'au fascicle et vingt dans 1'ambulacre posterieur. La f ossette posterieure gauche 
ne represente pas une depression continue et unique : elle est plutot formee par deux 
depressions successives separees par une region moins enfoncee ; cette disposition est 
evidemment accidentelle car elle n'existe pas du cote droit. 

Bien que les poches incubatrices soient moins developpees dans cet exemplaire 
que dans le precedent, elles existent cependant et il ne me parait pas douteux que 
1'exemplaire ne soit une femelle ; la disposition des ambulacres dorsaux est en effet 
bien differente chez le male qu'il me reste a decrire. 

J'ai etudie les caracteres du male sur deux exemplaires de tailles un peu differentes, 
et dont les dimensions respectives sans les piquants sont les suivantes : 

Longueur 34 millim. 29-5 millim. 

Largeur maxima .... 30 ,, 26 

Hauteur maxima .... 15 ,, 18'5 ,, 

Je represente ici le plus grand individu (PI. IV, fig. 3 a 5). 

Le test est un peu plus elargi et plus court que chez la femelle qui m'a 
servi de type pour la description de 1'espece, mais ce caractere ne doit pas etre lie a 
une difference de sexe, ainsi que nous 1'avons Vu plus haut. 

Dans les deux individus, 1'ambulacre anterieur dorsal est a peine deprime et il 
arrive a fleur du test en atteignant le bord interne du fascicle. Les quinze premieres 
paires de pores seules sont assez developpees, et les suivantes sont a peine apparentes ; 
les pores sont relativement un peu plus gros dans le plus petit exemplaire que dans 
1'autre. Les ambulacres lateraux anterieurs et posterieurs sont faiblement deprimes ; 
les ambulacres anterieurs sont un peu plus enfonces que les posterieurs, mais la 
difference est peu sensible. La depression commence des la premiere paire de pores 
dans les ambulacres posterieurs et seulement a partir de la deuxieme ou de la troisieme 

BRIT. ANTAECT. EXPED. 1907-9. VOL. II. I 



58 R. KCEHLER 

paire dans les anterieurs. Je compte vingt paires de pores dans chacun de ces 
ambulacres. 

Les orifices genitaux sont petits. Dans le grand individu, les deux orifices gauches 
sont tres allonges, presque deux fois plus longs que larges,et leur grand axe est oriente 
suivant la direction de 1'interradius : ils sont par consequent diriges obliquement 1'un 
par rapport a 1'autre. L'orifice droit est au contraire presque circulaire et plus petit 
que les deux autres ; le bord interne de cet orifice est separe du bord interne de 
1'orifice posterieur gauche par une distance de 1'7 millim. 

Dans le petit exemplaire, les trois orifices genitaux sont petits et circulaires, et 
1'orifice droit est plus ecarte de 1'orifice gauche que sur 1'autre individu, car il en 
est separe par un intervalle de 2 millim. La plaque madreporique n'offre qu'une 
petite plage perforce arrondie, autour de laquelle se montrent quelques tubercules, 
tandis que dans le grand individu, cette plaque presente la structure que j'ai decrite 
plus haut chez la femelle. 

Le fascicle offre aussi le meme trajet que chez cette derniere, mais, dans le petit 
echantillon, ses sinuosites sont a peine marquees. 

Le periprocte est beaucoup plus haut que large : il mesure 4' 7 millim. de hauteur 
sur 3'5 de largeur dans le grand exemplaire et 4 '4 sur 3 millim. dans le petit ; il est 
done comparativement un peu plus haut chez ce dernier. 

Je prie Sir Ernest Shackleton, commandant de 1'Expedition Antarctique Anglaise, 
de vouloir bien accepter la dedicace de cette espece. 

Rapports el differences. Mortensen a precise recemment les limites respectives des 
genres Hemiaster et Abatus et il a fait une revision des especes appartenant a ce dernier, 
especes qui sont toutes australes. En raison de la constitution de son appareil apical, 
VA. shackletoni appartient aux Eihmolysii de Loven, et, par consequent, il doit etre 
range dans le genre Abatus tel que Mortensen 1'admet. Le savant naturaliste danois 
a rendu aux zoologistes le tres grande service de fixer les caracteres des differentes 
especes du genre Abatus, especes qui avaient ete plus ou moins confondues par les 
auteurs en raison des descriptions insufnsantes qui en avaient ete donnees. Ses 
recherches 1'ont conduit a considerer, comme distinctes, les quatre especes suivantes : 

Abatus cordatus (Verrill). 
Abatus agassizii (Pfeffer). 
Abatus cavernosus (Philippi). 
Abatus philippii Loven. 

Quant a VA. elongatus que j'ai decrit sous le nom d' Hemiaster elongatus, Mortensen 
incline a le reunir a VA. agassizii. 

"L'A. cordatus provient des iles Kerguelen ; les trois especes suivantes appartien- 
nent toutes trois a rextremite meridionale de rAmerique du Sud et VA. elongatus 
a ete rencontre aux Orcades du Sud, 



ECHINODERMA 59 

L'A. shackletoni est depourvu de tubes ambulacraires sous-anaux, caractere qu'il 
partage avec les A. cordatus et agassizii ; il se distingue ainsi immediatement des 
A. philippii et cavernosus qui ont, de plus, le sillon dorsal tres marque et il s'ecarte 
surtout de \'A. philippii par les quatre petales dorsaux profondement deprimes et 
formant des poches incubatrices egalement developpees. 

Les caracteres de \'A. cordatus ont etc bien etablis par Mortensen : cette espece est 
tout a fait speciale aux iles de Kerguelen et elle ne penetre pas dans la region sud- 
americaine. ISA. shackletoni s'en distingue facilement par la presence de pedicellaires 
globiferes, par le sillon anterieur dorsal moins enfonce, par les poches incubatrices 
plus etroites et comparative ment plus longues, par une forme differente du corps qui 
est plus allonge et par 1'appareil apical qui est reporte en avant. 

La position de 1'appareil apical permet aussi de separer 1'A. shackletoni de 
YA. agassizii chez lequel cet appareil est central ; le test est plus allonge et le peri- 
stome est plus petit dans la premiere espece, enfin les pedicellaires sont differente : 
je n'observe pas, sur la face interne des valves des pedicellaires tridactyles de 
YA. shackletoni, cette saillie mediane formee par les mailles du reseau que Mortensen 
a indiquee chez YA. agassizii. 

La forme allongee de YA. shackletoni rappelle celle de YA. elongalus. Laissant de 
cote la question posee par Mortensen, de savoir si YA. elongatus doit etre reuni ou non 
a YA. agassizii, il est certain que YA. shackletoni ne peut etre confondu avec la 
premiere espece : il s'en eloigne par son appareil apical reporte en avant, par le 
periprocte plus grand et par une forme differente des pedicellaires tridactyles et 
rostres. 

Les spicules des tubes ambulacraires ne presentent aucun caractere particulier : 
ce sont des batonnets incurves avec des dents laterales inegales qui peuvent s'allonger 
et se reunir pour constituer de petites plaques perforees irregulieres. 

Je suis heureux de remercier ici mon excellent ami, M. le Dr. Th. Mortensen, de 
1'examen qu'il a bien voulu faire de cet Abatus ainsi que du genre que je decris 
ci-dessous ; il a eu egalement 1'extreme obligeance de me communiquer les epreuves 
du texte et des planches de son memoire, encore sous presse lorsque j'ecrivais ces lignes : 
Echinoidea of the Swedish South-polar Expedition, dans lequel il etudie les Spatangides 
antarctiques et qui m'a rendu les plus grands services pour mes comparaisons. 

Aux echantillons que j'ai indiques plus haut, il y a lieu d'aj outer encore cinq 
exemplaires extremement jeunes, dont la longueur varie entre 5 et 12 millim. et que 
je rapporte egalement a YA. shackletoni. Ces exemplaires etaient associes a six autres 
Oursins Irreguliers de dimensions analogues et que je considere comme appartenant 
au genre Brisaster : ce sont peut-etre des B. antarcticus Sladen, mais ils sont trop 
jeunes pour etre determines avec certitude et ils ne m'ont presente que quelques 
pedicellaires non caracteristiques : je ne les signale ici que pour memoire. 



60 R. KCEHLER 

PSEUDABATUS, HOV. geil. 

Ce genre est voisin du genre Abatus qu'il rappelle par la constitution de son appareil 
apical dans lequel la plaque madreporique separe largement les deux plaques genitales 
posterieures, mais il en diflere par la situation des poches incubatrices. Tandis que 
dans le genre Abatus, celles-ci commencent a 1'appareil apical ou a une tres petite 
distance de ce dernier, ici les poches ne se forment qu'a une certaine distance de 
1'appareil apical, apres la huitieme ou la neuvieme paire de pores ; il reste done, entre 
1'appareil apical et chacune des poches incubatrices, une partie des ambulacres non 
modifiee et tout a fait a fleur du test, tandis que la depression qui forme la poche se fait 
tres brusquement; le bord externe decelle-ci est voisin du fascicle. L'ambulacre anterieur 
reste aussi a fleur du test. Les orifices genitaux sont au nombre de trois. La face 
posterieure n'est pas verticale, mais elle est dirigee obliquement en avant du cote 
ventral, de telle sorte que le periprocte, qu'elle porte, est subventral. Dans la seule 
espece connue, les pedicellaires appartiennent aux trois types didactyle, rostre et 
globifere. Les pedicellaires tridactyles font completement defaut et les pedicellaires 
globiferes sont d'une forme particuliere, leurs valves ayant I'extiemite arrondie et 
munie de petites dents. 

Bien que le genre Pseudabatus soit voisin du genre Abatus, il m'a paru necessaire 
de 1'en separer en raison des trois caracteres principaux indiques ci-dessus : position 
des poches incubatrices eloignees de 1'appareil apical, situation du periprocte et forme 
des pedicellaires. 

PSEUDABATUS NIMRODI, nov. sp. (PL VII, fig. 1 a 8 ; PL VIII, fig. 7 a 12) 
Baie du Cap Royds. Juin 1908. Profondeur, 7-20 brasses. Deux echantillons. 

Les deux exemplaires ne sont pas en parfait etat d'integrite et ils presentent des 
cassures, soit sur la face ventrale, soit sur les cotes du test ; ils sont neanmoins tres 
suffisants pour permettre une etude complete. 

Ils ont tous deux a peu pres les memes dimensions qui sont les suivantes : longueur, 
49 millim. ; largeur, 43 millim. ; hauteur, 27 millim. 

Dans 1'un des individus, que j'appellerai 1'individu A, et qui est represente PL VII, 
fig. 1, et PL VIII, fig. 8, 10 et 12, la face dorsale du test offre, quand on regarde 
1'oursin de cote (PL VII, fig. 1), une courbe reguliere aussi bien en avant qu'en arriere 
de 1'appareil apical, de telle sorte que le contour de cette face est regulierement arrondi. 
L'autre individu, ou individu B (PL VIII, fig. 9), offre, dans la partie anterieure de 
la face dorsale, un profil simplement oblique et peu convexe, et la region interradiale 
posterieure est peu proeminente, de telle sorte que le test, vu lateralement, est plus 
conique que dans 1'autre exemplaire. Chez tous deux d'ailleurs, le pole apical est 
aplati. 



ECHINODERMA 61 

Vu par la face dorsale (PI. VIII, fig. 7 et 8), le contour de la face dorsale est ovalaire 
mais il n'est parfaitement regulier chez aucun des deux echantillons, et le cote droit 
du bord anterieur du test est un peu plus preeminent que le cote gauche. Le bord 
anterieur n'offre pas la moindre trace d'echancrure en son milieu car I'ambulacre 
anterieur est a fleur du test. 

La face ventrale est presque plane, et 1'interradius impair posterieur est tres peu 
convexe. Le bord anterieur du labre se trouve a 15 '5 millim. en arriere du bord 
anterieur du test chez 1'individu A ; la mesure n'a pas pu etre prise sur 1'individu B, 
dont la face ventrale est brisee dans la partie centrale. 

En examinant 1'echantillon A de profil, on constate que le bord superieur de la 
face dorsale se relie a la face ventrale par une petite face posterieure tres courte et 
oblique ; dans I'echantillon B, cette face est un peu plus haute et moins oblique. II 
en resulte que dans le premier, le periprocte est en partie place sur la face ventrale et 
il est visible quand on regarde 1'oursin par cette face (PI. VIII, fig. 12) ; dans le second, 
il n'apparait que fort peu du cote ventral. D'ailleurs dans 1'individu A, le periprocte 
est assez petit : il ne mesure que 5 millim. de diametre et il est a peu pres aussi haut 
que large ; son contour est irregulierement circulaire avec une petite pointe du cote 
ventral (PI. VIII, fig. 10). Dans 1'individu B, le periprocte est plus grand : il mesure 
6 '5 millim. de largeur sur 6 de hauteur ; son contour est aussi irregulierement arrondi 
(PI. VIII, fig. 11). 

Le pole apical est plus rapproche du bord anterieur du test que du bord posterieur. 
La ligne qui reunit les centres des deux orifices genitaux posterieurs se trouve a 
20 millim. du bord anterieur et a 29 millim. du bord posterieur du test. Les trois 
orifices genitaux de 1'exemplaire A sont legerement ovoiides et un peu plus grands 
que dans le second : ils mesurent 1*7 millim. de largeur et les centres des deux orifices 
anterieurs sont separes par un intervalle de 6 millim. (fig. 8) ; dans 1'exemplaire B, 
ces orifices ont 1'2 millim. de largeur, et les centres sont separes par une distance de 
5 millim. (fig. 7). 

Les contours des plaques de 1'appareil apical se reconnaissent facilement, surtout 
dans I'echantillon B (PI. VII, fig. 4). Dans ce dernier, les deux plaques genitales gauches 
sont une fois et demie plus longues que larges ; 1'orifice que chacune d'elles porte est 
beaucoup plus rapproche du bord externe que du bord interne, et il existe un certain 
nombre de petits tubercules en dehors et en dedans de 1'orifice. Ces plaques sont separees, 
sur la moitie de leur longueur, par la plaque ocellaire de I'ambulacre anterieur gauche. 
La plaque genitale droite est plus petite que les deux autres et a peine plus longue 
que large ; 1'orifice se trouve place a peu pres a egale distance du bord externe et du 
bord interne. La plaque madreporique, qui separe les deux plaques genitales gauches 
de la plaque droite, est allongee et retrecie dans sa region anterieure. Elle depasse en 
avant le milieu de la plaque ocellaire anterieure, et elle s'arrete, en arriere, au 
niveau du milieu des deux plaques ocellaires posterieures. Elle offre des orifices, 
petits et peu nombreux, irregulierement disposes et se montrant surtout dans sa^ 



62 R. KCEHLER 

region moyenne, tandis que sur le reste de sa surface, elle porte de petits tubercules. 
Les plaques ocellaires sont triangulaires, avec un angle interne arrondi et un bord 
externe plus ou moins excave, vers le milieu duquel se trouve le pore qui est 
allonge dans le sens radiaire ; ces plaques sont plus saillantes que les ambulacres 
qu'elles terminent. La plaque ocellaire de 1'ambulacre IV, qui est placee entre les 
moities externes des deux plaques genitales gauches, a Tangle proximal plus aigu 
et moins arrondi que les autres ; c'est la seule qui ne soit pas contigue a la plaque 
madreporique. 

Chez 1'individu A (PI. VIII, fig. 8), je constate, dans les caracteres de 1'appareil apical, 
quelques differences qui ne tiennent pas a une difference de sexe, car j'ai reconnu chez tous 
deux la presence d'oeufs dans les poches incubatrices. Les orifices genitaux sont un 
peu plus grands et plus ecartes que dans 1'echantillon B et ils sont aussi legerement 
ovalaires; les plaques genitales portent des tubercules plus nombreux. Les plaques 
ocellaires ne sont pas saillantes ; elles sont plus etroites et relativement un peu plus 
allongees que dans 1'individu B, et leurs contours sont moins distincts. La plaque 
madreporique ne presente de pores que sur une plage centrale peu etendue, et tout le 
reste de sa surface est couvert de petits tubercules secondaires. 

L'ambulacre anterieur impair reste absolument a fleur du test dans 1'individu A 
(PI. VIII, fig. 8) et il offre, dans sa moitie proximale, une tres legere tendance a se deprimer 
chez le second individu (fig. 7). Les zones poriferes sont droites et paralleles, et chacune 
d'elles renferme vingt-deux paires de pores comptes jusqu'au fasciole. Les pores des 
trois ou quatre premieres paires sont tres fins, tres rapproches et disposes obliquement 
par rapport a 1'axe de 1'ambulacre ; les suivants sont plus grands, plus ecartes et 
places transversalement. A partir de la dix-septieme ou de la dix-huitieme paire, les 
pores se rapprochent de nouveau en se placant obliquement et ils deviennent tres 
petits. Les pores sont voisins des bords anterieur et externe de la plaque ; quelques 
petits tubercules tres fins se montrent, soit entre le bord interne, soit entre le bord 
posterieur de la plaque et la paire de pores ; parmi les tubercules internes, on en 
remarque souvent un plus gros que les autres. A mesure qu'on se rapproche du 
fasciole, le nombre et la taille des tubercules augmentent et chaque plaque arrive a 
porter quelques petits tubercules primaires. Au-dela du fasciole, ces tubercules 
primaires deviennent plus nombreux et plus serres,et,dans leurs intervalles,setrouvent 
des tubercules beaucoup plus fins. 

Les ambulacres lateraux anterieurs sont tres forte ment divergents ; au contraire 
les ambulacres posterieurs sont tres rapproches. L'ambulacre anterieur forme, avec 
1'ambulacre posterieur du meme cote un angle a peu pres droit. 

L'ambulacre anterieur debute par une partie amincie qui va en s'elargissant et qui 

reste absolument au niveau du test, sans la moindre tendance a s'enfoncer, cela jusqu'a 

la neuvieme paire de pores. A ce niveau, 1'ambulacre se deprime brusquement en 

meme temps qu'il s'elargit, et il ne suit pas, en s'enfongant, une direction verticale 

. seulement, mais il se dirige obliquement en dedans vers le pole apical, de telle sorte 



ECHINODERMA 63 

que la poche incubatrice ainsi constitute est en realite plus longue qu'elle ne le parait 
exterieure ment, et que son bord interne, aminci, surplombe une portion de cavite 
qui peut atteindre 3 millim.de longueur environ. La cavite incubatrice atteint les 
dimensions suivantes : 

Echantillon A. Echantillon B. 

Longueur exterieure ... 11 millim. . 14 millim. 

Largeur exterieure . . . 5'5-, r . 6 ,, 

Profondeur maxima . . . 6 ,, . 8 '9 

Vers son tiers externe, la poche se retrecit et son fond se releve progressivement pour 
revenir au niveau du test ; elle se termine a peu pres au bord interne du fascicle dans 
1'echantillon B, tandis que dans 1'autre elle n'atteint pas exactement ce bord : il 
reste au moins a fleur du test une plaque ambulacraire qui separe la poche du fascicle 
et le relevement de cette poche est plus progressif. 

Les ambulacres posterieurs presentent des dispositions analogues. La poche, qui 
se forme brusquement, et dont la face interne est egalement dirigee obliquement en 
dedans, est separee de 1'appareil apical par une partie ambulacraire non modifiee et 
restant a fleur du test, sur laquelle je compte neuf paires de pores dans 1'individu B 
et huit dans 1'autre ; dans le premier, la poche est un peu plus grande. Les poches 
ont les dimensions respectives suivantes : 

Echantillon A. Echantillon B, 

Longueur exterieure . . 12 millim. . 14 millim. 

Largeur exterieure ... 6 ,, . 6 ,, 

Profondeur maxima . . . 9 ,, . 9 ,, 

Les poches posterieures sont done un peu plus grandes que les anterieures ; dans 
1'echuntillon B, elles s'etendent jusqu'au bord interne du fascicle, tandis que dans 
1'autre individu, elles sont separees de ce bord par une ou deux plaques a fleur 
du test, comme cela arrive pour les poches anterieures. Les plaques qui ferment 
les parois des poches presentent de petits tubercules portant de petits piquants 
fins et cylindriques, au milieu desquels j'ai trouve quelques oeufs chez les deux 
individus. 

Au dela des poches, les plaques ambulacraires sont uniformement couvertes de 
tubercules primaires entremeles d'autres tubercules beaucoup plus fins. 

Les regions interradiales de la face dorsale off rent egalement un recouvrement 
uniforme de tubercules primaires rapproches, entre lesquels se montrent des 
granules tres fins. L'interradius posterieur est simplement convexe sans former de 
gibbosite. 

La face ventrale est tres peu convexe (PL VIII, fig. 12). Le peristome est assez 
large. Le bord anterieur du labre est saillant et large, mais il est arrondi et pro- 
emine peu en avant ; il laisse a decouvert la plus grande partie des plaques du 
peristome, II m'a ete absolument impossible de reconnaitre, sur 1'exemplaire ou 



64 R. KCEHLER 

il est conserve, les limites posterieures du labre, meme apres traitement a 1'eau de 
Javelle. 

L'ambulacre anterieur ventral est legerement deprime ; il est assez large au 
voisinage du peristome, puis il se retrecit et se releve en se rapprochant du bord 
anterieur du test. II offre, de chaque cote, cinq pores ambulacraires places chacun 
au bord d'une depression ovalaire bien marquee. Les ambulacres lateraux anterieurs 
sont presque situes sur le prolongement 1'un de 1'autre. Us sont deprimes dans leur 
region proximale, un peu plus meme que 1'ambulacre anterieur, mais ils reviennent au 
niveau du test vers le milieu de leur longueur. Chacun d'eux oflre, de chaque cote, cinq 
ou six pores avec leurs depressions ovalaires. 

Les avenues ambulacraires posterieures sont larges et elles ne sont un peu 
deprimees qu'au voisinage immediat du peristome, ou chacune d'elles offre deux 
rangees legerement divergentes de quatre ou cinq pores avec leurs depressions ovalaires. 
Les plaques suivantes, tres grandes, presentent comme d'habitude, un petit pore place 
vers le bord anterieur. 

Les ambulacres n'offrent que quelques tubercules tres fins, peu serres sur les 
ambulacres anterieurs, plus serres sur les avenues ambulacraires posterieures. Les 
tubercules primaires n'y font leur apparition qu'au voisinage du bord du test. Les 
regions interradiales ventrales possedent des tubercules primaires sensiblement plus 
gros que ceux de la face dorsale, avec des tubercules tres fins qui forment generalement 
un cercle autour des precedents. Les tubercules de 1'interradius posterieur ne sont 
pas plus developpes que les autres : ils forment des files plus ou moins regulieres et 
divergentes partant de Tangle posterieur, qui debutent par des tubercules d'abord plus 
petits et qui grossissent ensuite a mesure qu'on se rapproche des avenues ambulacraires 
ventrales ou du peristome. 

J'ai dit plus haut que dans 1'echantillon A, 1'extremite posterieure etait legerement 
arrondie, tandis que dans le second, cette extremite etait mieux marquee et formait 
une petite face dirigee obliquement vers le bas ; aussi le periprocte est-il plus visible 
par la face ventrale dans 1'echantillon A que dans 1'echantillon B, bien qu'il soit plus 
petit dans le premier. Les plaques qui recouvrent le periprocte sont irregulieres 
comme forme et comme disposition, et elles sont plus grandes vers la peripherie. 

II n'existe pas de traces de tubes ambulacraires sous-anaux. 

Le fasciole se trouve tres rapproche du bord anterieur du test dont il est separe 
a peine par une distance de 5 millim., mais, comme la face dorsale s'inflechit tres 
rapidement en avant pour rejoindre la face ventrale et se reunit finalement a cette 
face par une partie a peu pres verticale, comme d'autre part le fasciole se trouve place 
sur cette partie presque verticale, celui-ci est a peine visible quand on regarde 1'animal par 
en haut. Apres un court trajet parallelement au bord du test, le fasciole s'en ecarte 
legerement et il offre deux concavites peu profondes separees par un angle aigu. II 
atteint ensuite le bord externe de la poche incubatrice anterieure qu'il contourne et 
s'inflechit d'abord en dedans, parallelement au bord posterieur de cette poche. II ne 



ECHINODERMA 65 

tarde pas, en formant un angle obtus, a reprendre son trajet parallele au bord du test 
dont il se trouve assez ecarte entre les deux poches incubatrices, puis, arrive a 7 ou 
8 millim. de la poche posterieure. il forme un nouvel angle obtus, se rapproche du bord 
du test, contourne le bord externe de la poche incubatrice posterieure et se dirige vers 
la ligne interradiale mediane en restant a 8 millim. environ du bord posterieur du test. 
Sa largeur moyenne est d'un millimetre environ ; il s'amincit legerement dans ses 
parties anterieure et posterieure et s'elargit -nn- peu au niveau des deux poches 
incubatrices. 

Les tubercules primaires ne sont pas tres saillants. Us sont plus gros sur la face 
ventrale que sur la face dorsale ainsi que j'ai eu 1'occasion de le dire, mais leurs 
dimensions restent tres uniformes dans la meme region ; ils deviennent seulement un 
peu plus gros sur les bords anterieur et posterieur des poches incubatrices. Ils sont 
performs et finement creneles. 

Les piquants que portent les tubercules primaires de la face dorsale sont tres 
courts, fins et serres, separes par d'autres piquants beaucoup plus petits. Ils sont un 
peu aplatis et elargis en spatule, avec 1'extremite tronquee et leur surface est finement 
striee ; ils deviennent un peu plus forts a 1'ambitus. Vers les poches incubatrices^ 
les piquants deviennent plus larges et plus forts, leur extremite est plus elargie et ils 
se dirigent horizontalement en s'enchevetrant avec leurs congeneres du bord oppose 
de maniere a recouvrir la poche. Ces piquants, ainsi que les piquants marginaux, 
offrent souvent une cannelure longitudinale assez marquee. Les piquants de la face 
ventrale sont plus developpes que sur la face dorsale ; ils sont allonges, aplatis, elargis- 
a 1'extremite et les plus gros sont canneles. 

Les pedicellaires appartiennent aux trois types didactyle, rostre et globifere. Les 
pedicellaires didactyles sont abondamment repandus sur tout le test, (PI. VII, fig. 6 
et 7). Leur pedoncule est relativement assez long et la tige calcaire est eloignee 
de la tete d'une distance qui est presque egale a la hauteur de celle-ci, qui varie entre 
0'6 et 0'7 millim. Les valves sont apiaties, et leur base offre une echancrure profor.de ; 
leur face externe est convexe, et, en dedans, elles s'adossent a leur congenere par 
un bord droit muni de denticulations extremement fines et tres regulieres. Les per- 
forations sont disposees regulierement. Les pedicellaires tridactyles, que ceux-ci 
remplacent, font completement defaut. 

Les pedicellaires rostres sont tres abondants aussi (PI. VII, fig. 8). Leur tete 
mesure 0'8 a 1 millim. de hauteur dans les plus grands et la tige calcaire de leur 
pedoncule, elargie a son extremite, en est tres rapprochee. Leurs valves offrent une 
base allongee ; le limbe est etroit et il conserve a peu pres la meme largeur jusqu'a 
1'extremite qui est un peu elargie ; les bords sont reployes en dedans : ils sont un 
peu irreguliers, sinueux et ils ofErent quelques dents tres petites et basses, tres 
espacees, tandis que 1'extremite porte une rangee de dents tres fines, rapprochees et 
ties regulierement disposees. 

Les pedicellaires globiferes sont tres rares (PI. VII, fig. 5). Leur tete, qui mesure 

BEIT. ANTARCT. EXPED. 1907-9. VOL. II. K 



66 R. KCEHLER 

0'4 a 0'5 de hauteur, est recouverte d'un tegument glandulaire et fonce semblable 
a celui qu'on rencontre dans le genre Abatus. Les valves sont formees d'un tissu 
calcaire delicat avec de grandes perforations ; elles oflrent un limbe allonge, assez 
large et allant en s'elargissant legerement jusqu'a 1'extremite : celle-ci est arrondie 
et porte dix a douze dents coniques, pointues, assez courtes, contigues, un peu inegales 
et formant une bordure tout le long du bord terminal. L'orifice, tres large, 
depasse le milieu de la longueur du limbe, et, en dessous de lui, ce dernier constitue 
un tube dont les parois offrent de grandes perforations. 

La couleur des deux echantillons etait a peu pres uniformement noire. Pour les 
-etudier et les photographier, j'ai du les decolorer incompletement a 1'eau de Javelle, 
mais le test a conserve des taches et des marbrures: aussi j'ai eprouve de grandes 
difficultes pdur obtenir des photographies passables. 

LYON, Septembre 1910. 



LISTE DES OUVKAGES CITE'S 

'91. PERKIER, ED., Echinodermes de la Mission du Cap Horn. Paris, 1891. 

95. LKITPOLD, FB, " Asteroidea der Vettor Pis&ni Expedition " Zeit.fiir wiss. Zool., Bd. LIX. 

'03. LUDWIG, H. . Resultats du Voyage du S. 7. " Bdgica." Seesterne. 

05. i, " Asterien und Ophiuren der schwedischen Expedition nach den Magalhaenslandern, 

1895-97." Zeit. fur wiss. Zool., Bd. 82. 

06. FiSHER,3V. K. The Starfishes of the Hawa'ian Islands. U.S. Fish Commission Bulletin for 1903, 

Part III. Washington, 1906. 
06. KCEHLER, B. . Expedition antarctique framfaise commandee par le Dr. J. Charcot. Echinodermes. 

08. i, . " Asteries, Ophiures et Eehinides de 1'Expedition Antarctique Nationale Ecossaise." 

Trans. Roy. Soc. Edinburgh, vol. xlvi. 

08. BELL, J. . " National Antarctic Expedition," Natural History, vol. iv, : Echinoderma. 

10. MORTENSEX, Tl). " Die Echinoiden der deutschen siidpolar Expedition, 1901-03," Deutsch* sudpolar 

Expedition, xi. : Zoologie, iii. 
,10b:s. South-polar Expedition, vol. iii.: Ethinoidea. 



PLATE IV 



PLANCHE IV 

FIG..I. Cryaster antarcticus. Face dorsale reduite d'un cinquieme environ. 

FIG. 2. Cryaster antarcticus. Face ventrale reduite d'un cinquieme environ. 

FIG. 3. Abatus shackletoni, male. Vue laterale. Grossissement T6. 

FIG. 4 Abatus sJiacMetoni, male. Face posterieure. Grossissement l - 6 

FIG. 5. Abatus shackletoni, male. Face dorsale. Grossissement T6. 

FIG. 6. -Abatus shackletoni, femelle. Face posterieure. Grossissement To. 

FIG. 7. Abatus shackletoni. Valve d'un pedicellaire tridactyle a valves larges vue 

de profil. Grossissement 60. 

FIG. 8. Abatus sliackletoni. Meme valve vue de trois quarts. Grossissement 60. 
FIG. 9. Abatus shackletoni. Valve d'un pedicellaire rostre vue par la face externe. 

Grossissement 80. 
FIG. 10. Abatus shackletoni. Valve d'un pedicellaire rostre vue par la face interne. 

Grossissement 80. 

FIG. 11. Abatus shackletoni. Valve d'un pedicellaire globifere. Grossissement 110. 
FIG. 12. Abatus shackletoni. Valve d'un pedicellaire tridactyle a valves etroites. 

Grossissement 60. 



BRIT. ANTARCT. EXPED. 1907-9 



VOL. II. PLATE IV 



11. KCEHLER: ECHINODERMES 




R. Kd-liler fecit 



PLATE V 



PLANCHE V 

FIG. 1. Ophioglypha fiexibilis. Face dorsale. Grossissement 4. 

Fia. 2. Ophioglypha flexibilis. Face ventrale. Grossissement 4. 

FIG. 3. Coscinasterias victories. Face dorsale. Grossissement 1*2. 

FIG. 4. Coscinasterias victoria. Face ventrale. Grossissement 1*2. 

FIG. 5. Coscinasterias brucei. Jeune individu vu par la face dorsale. Grossisse- 
ment 1'8. 

FIG. 6. Notasterias armata. Pedicellaire macrocephale vu de profil. Grossisse- 
ment 25. 

FIG. 7. Notasterias armata. Pedicellaire macrocephale vu de profil apres traitement 
par la potasse qui a separe les deux valves et la piece basilaire. Grossisse- 
ment 25. 

FIG. 8. Notasterias armata. Valve isolee d'un petit pedicellaire macrocephale. 
Grossissement 40. 

FIG. 9. Notasterias armata. Deuxieme valve du meme pedicellaire. Grossissement 
40. 

FIG. 10. Notasterias armata. Piece basilaire du meme pedicellaire. Grossisse- 
ment 40. 

FIG. 11. Notasterias armata. Valve d'un autre pedicellaire macrocephale un peu 
plus gros que le precedent. Grossissement 30 



BRIT. ANTARCT. EXPED. 1907-9 



VOL. II. PLATE V 



11 KCEHLER: ECHIXODERMES 




R. Koehler fecit 



PLATE VI 



PLANCHE VI 

FIG. 1. Notasterias armata. Face ventrale du plus grand exemplaire. Grossisse- 
ment 2. 

FIG. 2. Notasterias armata. Face dorsale. Grossissement 2. 

FIG. 3. Notasterias armata. Pedicellaire droit du sillon ambulacraire . Grossisse- 
ment 35. 

FIG. 4. Notasterias armata. Gros pedicellaire macrocephale entier vu de face. 
Grossissement 20. 

FIG. 5 a 7. Notasterias armata. Les deux valves et la "piece basilaire d'un gros 
pedicellaire macrocephale. Grossissement 20. 

FIG. 8. Notasterias armata. Pedicellaire macrocephale de taille moyenne vu de face. 
Grossissement 35. 

FIG. Q. Ophiodiplax disjuncta. Echantillon recueilli par 1'Expedition Antarctique 
Anglaise ; face dorsale. Grossissement 4.. 

FIG. 10. Ophiodiplax disjuncta. Echantillon recueilli par la deuxieme Expedition 
Antarctique du Dr. Charcot ; face dorsale. Grossissement 4. 

FIG. 11. Ophiodiplax disjuncta. Face ventrale du meme echantillon. Grossisse- 
ment 4. 



BRIT. ANTABCT. EXPED. 1907-9 

R. KCEHLER: ECHINODERMES 



VOL. II. PLATE VI 




R. K<*>hler fecit 



PLATE VII 



BRIT. ANTARCT. EXPED. 1907-9. VOL. II. 



PLANCHE VII 

FIG. 1. Pseuddbatus nimrodi, individu A. Vue laterale. Grossissement 1'4. 

FIG. 2. Pseudabatus nimrodi, individu B. Vue laterale. Grossissement 1*4. 

FIG. 3. Pseudabatus nimrodi, individu B. Face ventrale. Grossissement 1*4. 

FIG. 4. Pseudabatus nimrodi, individu B. Region apicale. Grossissement 4. 

FIG. 5. Pseudabatus nimrodi. Valve d'un pedicellaire globifere. Grossissement 100. 

FIG. 6. Pseudabatus nimrodi. Valve d'un pedicellaire didactyle. Grossissement 65. 

FIG. 7. Pseuddbatus nimrodi. Pedicellaire didactyle. Grossissement 35. 

FIG. 8. Pseuddbatus nimrodi. Valve d'un pedicellaire rostre. Grossissement 55. 

FIG. 9. Ophioglypha resistens. Face dorsale. Grossissement 3 '2. 

FIG. 10. Ophioglypha resistens. Face ventrale. Grossissement 3 '2. 

FIG. 11. Ophioglypha resistens, jeune individu. Face dorsale. Grossissement 6. 

FIG. 12. Ophioglypha resistens. Face ventrale du meme individu. Grossissement 6. 

FIG. 13. Ophiodiplax disjuncta. Echantillon de FExpedition Antarctique Anglaise ; 

face ventrale. Grossissement 4. 

FIG. l4:. Amphiura algida. Face dorsale. Grossissement 14. 
FIG. 15. Amphiura algida. Face ventrale. Grossissement 14. 



BRIT. ANTARCT. EXPED. 1907-9 

R. KXEHLER: ECHINODERMES 



VOL. II. PLATE VII 




R. Ku-hlcr fecit 



PLATE VIII 



PLANCHE VIII 

FIG. 1. Abatus shackletoni, individu femelle muni de ses piquants. Face dorsale. 

Grossissement 1'6. 
FIG. 2. Abatus shackletoni, meme individu vu par la face ventrale. Grossissement 

re. 

FIG. 3. Abatus shackletoni, individu femelle en partie depouille de ses piquants. 

Face ventrale. Grossissement 1'6 

FIG. 4. Abatus shackletoni, meme individu. Vue laterale. Grossissement I 1 6. 
FIG. 5. Abatus shackletoni, meme individu. Face dorsale. Grossissement 1'6. 
FIG. 6. Abatus shackletoni. Face dorsale d'un autre individu femelle. Grossisse- 
ment 1'6 

FIG. 7. Pseudabatus nimrodi, individu B. Face dorsale. Grossissement 1'4. 
FIG. 8. Pseudabatus nimrodi, individu A. Face dorsale. Grossissement 1 ' 4. 
FIG. 9. Pseudabatus nimrodi, individu B. Vue laterale. Grossissement 1'4. 
FIG. 10. Pseudabatus nimrodi, individu A. Face posterieure. Grossissement 1*4. 
FIG. 11. Pseudabatus nimrodi, individu B. Face posterieure. Grossissement 1* 4. 
FIG. 12. Pseudabatus nimrodi, individu A. Face ventrale. Grossissement 1*4. 



BKIT. ANTARCT. EXPEI). 1907-9 



VOL. II. PLATE VIII 



K. KCEHLER: ECHINODERMES 




R. Kd-hlur fecit 



REPORTS IN PREPARATION 

FRESH-WATER ALG^. BY W. WEST, F.L.S., AND 

PROF. G. S. WEST, M.A., D.Sc. 

LICHENS. BY DR. O. V. DARBISHIRE, NEWCASTLE. 
AVES. BY .1. MURRAY. 
MAMMALIA. Bv J. MURRAY. 
PYCNOGONIDA. BY T. V. HODGSON. PLYMOUTH. 
ISOPODA. BY T. V. HODGSON, PLYMOUTH. 
AMPH1PODA. BY T. V. HODGSON, PLYMOUTH. 
CUMACEA. BY DR. W. T. CALMAN, BRITISH MUSEUM. 
VERMES. BY E. J. GODDARD, B.A., B.Sc., SYDNEY. 
ROTIFERA OF NEW ZEALAND, AUSTRALIA, PACIFIC ISLANDS 

AND CANADA. BY J. MURRAY. 

POLYZOA. BY W. D. HENDERSON, M.A., B.Sc., Pn.D., BRISTOL. 
ALCYONARIA. KY PROF. J. ARTHUR THOMSON, M.A., ABERDEEN. 
PORIFERA. BY R. KIRKPATRICK, BRITISH MUSEUM (Natural 

History). 

LIFE UNDER POLAR CONDITIONS. BY J. MURRAY. 
STUDIES IN GEOGRAPHICAL DISTRIBUTION. BY J. MURRAY. 
GENERAL GEOLOGY. BY PROF. T. W. E. DAVID, F.R.S., SYDNEY, 

AND R. E. PRIESTLEY. 

GLACIOLOGY. BY PROF. T. W. E. DAVID, F.R.S., SYDNEY. 
PETROLOGY AND MINERALOGY. BY DRS. MAWSON, WOOL- 

NOUGH, AND JENSEN. 

TIDES. BY Sm G. DARWIN, K.C.B., F.R.S., CAMBRIDGE. 
PHYSICS. BY DOUGLAS MAWSON, D.Sc., BE. 
MAGNETISM. BY COLERIDGE FARR, D.Sc., CHRISTCHL-RCH, N.Z. 
AURORA. BY DOUGLAS MAWSON, D.Sc., B.E., ADELAIDE. 
METEOROLOGY. BY H. A. HUNT, COMMONWEALTH METEOROLOGIST, 

ETC ETC. 



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