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PROCEEDINGS : |

AND

TRANSACTIONS

OF THT

LIVERPOOL BIOLOGICAL SOCIETY.

VOL. XXVI.

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CONTENTS.

I.—-PROcEEDINGS.

Office-bearers and Council, 1911-1912 . vu
Report of the Council . vill
Summary of Proceedings at the Meetings. 1x
List of Members . : ; : Xi

Treasurer’s Balance Sheet . A f ; é XV

I].—TRANSACTIONS.

Presidential Address—“ Reptiles in es: By
J. H. O’Connett, L.R.C.P. : 1
Twenty-fifth Annual Report of the Liverpool Marine
Biological Committee and their Biological Station
‘at Port Erin. By Prof. W. A. Herpmay, D.Sc.,
F.RB.S. ae : ; ot) els

Report on the Investigations carried on during 1911,
in connection with the Lancashire Sea-Fisheries
Laboratory, at the University of Liverpool, and
the Sea-Fish Hatchery at Piel, near Barrow ;
By Prof. W. A. Herpmay, D.Sc., F.R.S., ANDREW
Scott, A.L.S., and James Jounsrone, B.Sc. ae oa:

L.M.B.C. Memoir on “ Buccinum.” By W. Darin,
D.Sc. ; : : . : . 258

PROCEEDINGS

LIVERPOOL BIOLOGICAL SOCTETY

OFFICHE-BEARERS AND COUNCIL.
Gx- Presidents :

1886—87 Pror. W. MITCHELL BANKS, M.D., F.R.C.S.
1887—88 J. J. DRYSDALE, M.D.

1888—89 Pror. W. A. HERDMAN, D.Sc., F.R.S.E.
1889—90 Pror. W. A. HERDMAN, D.Sc., F.R.S.E.
1890—91 T. J. MOORE, C.M.Z.S.

1891—92 T. J. MOORE, C.M.Z.S.

1892—93 ALFRED O. WALKER, J.P., F.L.S.
1893—94 JOHN NEWTON, M.R. cs.

1894—95 Pror. F. GOTCH, MAS Ee. Be.

1895—96 Pror. R. J. HARVEY GIBSON, M.A.
1896—97 HENRY O. FORBES, LL.D., E.Z.8.
1897—98 ISAAC C. THOMPSON, F.L. S., F.R.M.S.
1898—99 Pror. C. S. SHERRINGTON, M.D. oy Hs
1899—1900 J. WIGLESWORTH, M.D., F.R.C.P.
1900—1901 Pror. PATERSON, M.D., M.R.C.S.
1901—1902 HENRY C. BEASLEY.

1902—1903 R. CATON, M.D., F.R.C.P.

1903—1904 Rev. T. S. LHA, M.A.

1904—1905 ALFRED LEICESTER.

1905—1906 JOSEPH LOMAS, F.G.S.

1906—1907 Pror. W. A. HERDMAN, D.Sc., F.R.S.
1907—1908 W. T. HAYDON, F.L.S.

1908—1909 Pror. B. MOORE, M.A., D.Sc.
1909—1910 R. NEWSTEAD, M.Sc., F.ES.
1910—1911 Pror. R. NEWSTEAD, M.Sc., F.R.S.

SESSION XXVI., 1911-1912.
President :

fate O'CONNELL, 1.8.C.P.

Vice- Presidents :
Pror. W. A. HERDMAN, D.Sc., F.R.S.
Pror. R. NEWSTEAD, M.S&c., F.B.S.

Hon. Creasurer : Hon. Hibrarian :
W. J. HALLS. MAY ALLEN, B.A.

Hon. Secretary:
JOSEPH A. CLUBB, D.Sc.

Council :
HENRY C. BEASLEY. Pror. B. MOORE, M.A., D.Sc.
H. CATON, M.D., F.R.C.P. DOUGLAS LAURIE, M.A.
H. B. FANTHAM, D.Sc., B.A. W.S. LAVEROCK, M.A., B.Sc.
Moet, HAYDON, F.L.S. Pror. SHERRINGTON, F.R.S.
J. JOHNSTONE, B.Sc. | W.M. TATTERSALL, D.Sc.
J. SHARE-JONES, F.R.C.V.S. K. THOMPSON.

Representative of Students’ Section :
R. ROBBINS (Miss),

Vili - YLIVERPOOL BIOLOGICAL SOCIETY.

REPORT of the COUNCIL.

Durine the Session 1911-12 there have been seven
ordinary meetings and one field meeting of the Society.

- The communications made to the Society at the
ordinary meetings have been representative of almost all
branches of Biology, and the various exhibitions and
demonstrations thereon have been of great interest.

Prof. F. W. Gamble, D.Sc., F.R.S., of Birmingham,
lectured before the Society, at the February Meeting,
on ‘‘ Methods and Results of Symbiosis.”’

The Library continues to make satisfactory progress,
and additional important exchanges have been arranged.

The Treasurer’s statement and balance-sheet are

appended.

The members at present on the roll are as follows :—
Ordinary members” - - - - - - 45
Associate members - - : - : - iS
Student members, including Students’ Section - 37

Total - eg

SUMMARY OF PROCEEDINGS AT MEETINGS. 1X

SUMMARY of PROCEEDINGS at the MEETINGS.

The first meeting of the twenty-sixth session was
held at the University, on Friday, October 183th, 1911.

The President-elect (J. H. O’Connell, L.R.C.P.)
took the chair in the Zoology Theatre.

1. The Report of the Council on the Session 1910-1911
(see “‘ Proceedings,’’ Vol: XXV., p. vill.) was
submitted and adopted.

The Treasurer’s Balance Sheet for the Session 1910-
1911 (see ‘‘ Proceedings,’ Vol. XXV, p. xx.)

was submitted and approved.

pwn

3. The following Office-bearers and Council for the
ensuing Session were elected :—Vice-Presidents,
tae Herdman, D-.Sc.; F.R:S:;:. and Prof.
Newstead, M.Sc., F.R.S.; Hon. Treasurer, W. J.
Halls; Hon. Librarian, May Allen, B.A.; Hon.
Secretary, Joseph A. Clubb, D.Se.; Council,
HeeC. Beasley, Dr. Caton, Dr. Fantham, W. T.
Haydon, F.L.8., J. Johnstone, B.Sc., J. Share-
tenes, F.R.C.V.8S., Prof. B. Moore, M.A., D.Sc.,
W. S. Laverock, M.A., B.Sc., Douglas Laurie,
Mem, Prof. Sherrington, F.R.S.; and E.
Thompson.

meee H. O'Connell, L.R.C.P., delivered the
Presidential Address on ‘‘ Reptiles in Captivity ”’
(see “‘ Transactions,’ p. 1). <A vote of thanks
was proposed by Dr. Dakin, seconded — by

Mr. Laurie, and carried with acclamation,

xX | LIVERPOOL BIOLOGICAL SOCIETY.

The second meeting of the twenty-sixth session was
held at the University, on Friday, November 10th, 1911.
The President in the chair.

1. Dr. Clubb exhibited with remarks a living specimen
of Comvys erosa, from West Africa.

2. Prof. Herdman exhibited a collection of Marine
Invertebrates dredged from Hebridean seas during
the summer. We 3

3. Prof. Herdman submitted the Annual Report on the
work of the Liverpool Marine Biology Committee
and the Port Hrin Biological Station (see
“Transactions, p13).

The third meeting of the twenty-sixth session was
held at the University, on Friday, December 15th, 1911.
The President in the chair.

1. Mr. H. C. Beasley exhibited with remarks some
specimens of peaty material from Leasowe, with
fungus investing it.

2. Dr. Dakin submitted a paper on the ‘‘ Osmotic
pressure of the blood of Aquatic Animals.’

The fourth meeting of the twenty-sixth session was
held at the University, on Friday, January 12th, 1912.
~The Vice-President (Prof. Herdman) in the chair.

1. Dr. H. B. Fantham communicated a paper on
| ‘“Some Flagellate Parasites of Insects—not
always harmful—and their relation to

SUMMARY OF PROCEEDINGS AT MEETINGS. X1

)

Trypanosomes.’’ The parasites discussed belong
to the genera Herpetomonas and Crithidia and
occur in the digestive tracts of insects such as
flies, mosquitos and water-bugs. The herpeto-
monad found in the latex of various Huphorbia

and the parasite of Kala-azar were mentioned.

The fifth meeting of the twenty-sixth session was
held at the University, on Friday, February 9th, 1912,

jointly with the Students’ Section of the Society.
1) Prof. F. W. Gamble, F.R.S., of Birmingham,

lectured before the Society on ‘* Methods and
Results of Symbiosis.”’

The sixth meeting of the twenty-sixth session was
held at the University, on Friday, March 8th, 1912.
The President in the chair.

1. Prof. Newstead lectured to the Society on ‘‘ Notes
on the Natural History of Nyassaland,’’ giving
a most interesting account of his recent visit to
that country.

The seventh meeting of the twenty-sixth session was
held at the University, on Friday, May 10th, 1912.
The President in the chair.

1. Prof. Herdman submitted the Annual Report of the

Investigations carried on during 191] in con-

X11 LIVERPOOL BIOLOGICAL SOCIETY.

nection with the Lancashire Sea Fisheries
Committee (see “‘ Transactions,’’ p. 71).

2. L.M.B.C. Memoir on the Whelk, by Dr. Dakin
(see “* Transactions,’’ p. 258).

The eighth meeting of the twenty-sixth session was
the Annual Field Meeting held at Huilbre Island, on
Saturday, June Ist. At the short business meeting held
after tea, on the motion of the President from the chair,
Mr. James Johnstone, B.Sc., was unanimously elected
President for the ensuing session.

LIST of MEMBERS of the LIVERPOOL
BIOLOGICAL SOCIETY.

SHSSLON 1911-1912.

A. OrpinARY MEMBERS.

ate (Life Members are marked with an asterisk.)
ELECTED.

Pee xbram, Prof. J. Hill,. 74, Rodney Street,
Liverpool.

1909 *Allen, Miss May, B.A., Hon. Lisrarran, Univer-
sity, Liverpool.

1910 Barratt, Dr. J. O. Wakelin, Cancer Research
Laboratory, University, Liverpool.

1888 Beasley, Henry C., 25a, Prince Alfred Road,
Wavertree.

0S) Bigland, H. D., 3B.A., Shrewsbury Road,
Birkenhead. |

1903 Booth, jun., Chas., 30, James Street, Liverpool.

1886 Caton, R., M.D., F.R.C.P., 78, Rodney Street.

1886 Clubb, J. A., D.Sc., Hon. Sucrerary, Free Public
Museums, Liverpool.

1909 Dakin, W., D.Sc., The University, Liverpool.

1911 Ellison, George, 4, Loudon Grove, Liverpool.

X1V

1910
1902
1886
1910
1896
1886
1893
1902
1903
1903

1898
1894

1896

1906
1905

1904
1904

1904
1903

1908

LIST OF MEMBERS.

Fantham, Dr. H. B., School of Tropical Medicine,
University, Liverpool.

Glynn, Dr. Ernest, 62, Rodney Street.

Halls, W. J., Hon. Treasurer, 35, Lord Street.

Hamilton, Mrs. J., 92, Huskisson Street, Liver-
pool. |

Haydon, W. T., F.L.S., 55, Grey Road, Walton,
Liverpool.

Herdman, Prof. W. A., D.Sc., F.R.S., Vucx-
PresipENT, University, Liverpool.

Herdman, Mrs. W. A., Croxteth Lodge, Ullet
Road, Liverpool.

Holt, sA:) Crofton; Arie burch:

Holt, George, Grove House, Knutsford.

Holt, Richard D., M.P., J, India) Sualdamee
Liverpool. :

Johnstone, James, B.Sc., University, Liverpool.

Lea, Rev. T. S., D.D., The Vicarage, St. Austell,
Cornwall.

Laverock, W. S., M.A., B.Sc., Free Museums,
Liverpool.

Laurie, R. Douglas, M.A., University, Liverpool.

Moore, Prof. B., D.Sc., F.R.8., -Wmiversiige
Liverpool.

Newstead, Prof. R., Vicr-Presipmnt, M.Sc.,
F.R.S., School of Tropical Medicine, Liverpool.

O’Connell, Dr. J. H., Prestpent, 38, Heathfield
Road, Liverpool.

Pallis, Miss M., Tatoi, Aigburth Drive, Liverpool.

Petrie, Sir Charles, Ivy Lodge, Ashfield Road,
Aigburth, Liverpool. ;

Rathbone, H. R., Oakwood, Aigburth.

1890 *Rathbone, Miss May, Backwood, Neston.

1910

1897
1908
1894
1908

1895

1886
1903
1903

1905
1889
1888
1891

L905
1905

1910
1903

1910

LIVERPOOL BIOLOGICAL SOCIETY. XV

Riddell, Wm., M.A., Zoological Department,
University, Liverpool.

Robinson, H. C., Malay States.

Rock, W. H., 25, Lord Street, Liverpool.

Scott, Andrew, A.L.S., Piel, Barrow-in-Furness.

Share-Jones, John, F.R.C.V.S., University,
Liverpool.

Sherrington, Prof., M.D., F.R.S., University,
Liverpool.

Smith, Andrew T., 21, Croxteth Road.

Stapledon, W. C., “ Annery,” Caldy, West Kirby.

Thomas, Dr. Thelwall, 84, Rodney Street, Liver-
pool. |

Thompson, Edwin, 25, Sefton Drive, Liverpool.

Thornely, Miss L. R., Nunclose, Grassendale.

Toll, J. M., 49, Newsham Drive, Liverpool.

Weaelesworth, J.. M.D., F.R.C.P., County

_ Asylum, Rainhill.

B. AssocrATE MEMBERS.

Carstairs, Miss, 39, Lilley Road, Fairfield.

Harrison, Oulton, Denehurst, Victoria Park,
Wavertree. ,

Kelley, Miss A. M., 10, Percy Street, Liverpool.

Tattersall, W. D., D.Sc., The Museum, Man-
chester.

Tozer, Miss IX. N., Physiology Laboratory, The

University, Liverpool.

XV1 LIST OF MEMBERS.

C. Uwniversiry STUDENTS’ SECTION.
President: Miss R.. Robbins.
Hon. Secretary: Miss C. M. P. Stafford.

Members :

The Misses Gleave, Robinson, JLatarche, Gill,

Kdmondson, Platt, Bradley, Cavanagh, Lewis,
Kay, E. Smith, [lhngworth, Thornton, Hewitt,
Hodgson, Quirk, Payne, Brew, Millican, Clerke,
Kirk, Udall, (Garside, Mitile; Higson, Robinson,
Clegg and Upson; Messrs. Waterhouse, Goodburn,
Rowlands, Barlow, Daniel and Hamilton.

D. Honorary MEMBERS.
S.A.S., Albert I., Prince de Monaco, 10, Avenue du

brocadéro, Paris.
Bornet, Dr. Edouard, Quai de la Tournelle 27, Paris.
Claus, Prof. Carl, University, Vienna.
Fritsch, Prof. Anton, Museum, Prague, Bohemia.
Haeckel, Prof. Dr. E., University, Jena.
Hanitsch, R., Ph.D., Raffles Museum, Singapore.
Solms-Laubach, Prof.-Dr., Botan. Instit., Strassburg.

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TRANSACTIONS

OF THE

LIVERPOOL BIOLOGICAL SOCIETY.

INAUGURAL ADDRESS
ON
REPTILE LIFE IN CAPTIVITY.

bye OuN FH. O’CONNELL,. L.R.C.P.,

PRESIDENT.

In selecting reptile life as the subject of this
evening’s address, I propose to deal with the matter from
the Natural History side of the question, and to show
how far it is possible to approximate their artificial to
their natural conditions. It is a frequent reproach
against this group that they are sluggish and more or
less inert, and in many cases this is so because the
conditions of their captivity are entirely unsuitable to
their lives. This group is not without its fascination,
and the romantic weirdness of its past history awakens
many an echo in the fairy tales of childhood and the

folk-lore of nations.

What an interesting ancestry to look back on, in the
dim and distant ages of the past, in the Jurassic period,
when vast tracts of primeval forests clothed the earth

with a plant life long since gone; the period of the

passing of the giant fern-like forms, and the great horse-
tails of the Carboniferous swamps, and the coming of the
Cycads. The glory of this race has gone into the long
and never-ending night where oblivion, in enshrouding
the past history of the race, has almost closed the book
of nature and, indeed, would have done so had they not
literally left ‘‘ foot prints in the sands of time’’; and
had not the casts of their bones been taken first in mud

and afterwards perpetuated in rock,

u TRANSACTIONS LIVERPOOL BioLOGicAL SOCIETY,

Our present interest is with the descendants of the
great past dead. The majority of reptiles come from
tropical countries, and this is the first difficulty in the
way of keeping them in a temperate climate. It is very
rare that the temperature of the British Isles is high
enough to suit these animals, and we must accordingly
resort to artificial heating. We must study the most
suitable temperature, light and feeding. However, there
are a few general considerations to be investigated first.
Most reptiles, when free, hibernate during the colder
months in their own climates, as also our own snakes and
lizards, but prior to doing so they store up sufficient fat
to carry them through their sleep. This raises an
interesting point as to whether hibernation is essential
to the well-being of a reptile or not; many foreign lizards
have lived more than one winter in our climate in an
artificial temperature, some indeed for several years,
without, apparently, any bad effects resulting from
suspension of hibernation. These cases would seem to
show that this resting stage is not a physiological
necessity to every lizard. On the other hand, some
years ago, I kept some green lizards which I had
collected in Jersey, partly through a winter at a
suitable heat. They did not do very well and most of
them died before the winter ended. I find the Amphibia
behave much as the reptiles do, for on another occasion
a British toad had been allowed to go into its winter
sleep and after two or three weeks it was put back into
‘the heated case. It was completely upset and did not
resume feeding for a very long time.

70° F. is a good average heat at which to maintain
most reptiles; this may be done by direct heat under
the case, the bottom of which is covered with sand, or
to have the case over a heated water bath. The best and

—Te. Tee © ei.
Cd nal

ae! ae
7: 6%

REPTILE LIFE IN CAPTIVITY. . 8

most economical method is by using a small copper boiler

with copper pipes, in which hot water can circulate in
a tray under the case.

Unfortunately, lizards require a considerable amount
of sunshine before they will feed or do well, and are
thus the most difficult group to deal with. Some of the
desert species are particularly difficult to keep.

_ The Crocodilia afte probably the easiest to manage.
Their cases should contain a tank sufficiently large to
enable them to move about with freedom, and deep
enough for them to remain completely submerged if they
so wish. In addition it is advisable to have a dry part
for them to come out of the water and bask in the sun.
They will feed freely on raw meat, fish, small frogs or
even insects. There is a marked difference in the
temperaments of these creatures. The long-snouted
W. African crocodile (C. cataphractus) is a timid animal,
not much given to biting, while its ally the common
crocodile (C. niloticus) is always vicious and treacherous :
C. americanus is very pretty when small and docile;
C. porosus, an Indian species, is usually vicious.
Osteolaemus tetraspis, from the W. Coast of Africa, is
an interesting species to keep; it is short and heavily built
and nicely mottled and marked.

Of all the Crocodilia, the Mississippi Alligator is
the most suitable for captivity as it is especially docile
and quickly recognises its feeder. The natural cry of
this group is something between a grunt and a bark, and
they are remarkably sensitive to the lightest touch over
the shields of their backs and sides, and will get into
the strangest positions in their endeavours to remove the
source of annoyance. Swimming is usually performed
by vigorous strokes of the tail with the limbs pressed
close to the sides of the body.

4 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

’ The water tortoises require similar conditions to
those of crocodiles. These may be easily distinguished
by the flattened webbed feet and* usually depressed
carapaces. Most of the water forms are animal feeders
and tear their food with their long and sharp claws.
It is necessary to be very careful in handling the larger
ones as they bite very badly, usually taking out a piece
of one’s finger.

The large Amazon fresh water tortoise (Podocnemis)
seems to be a vegetable feeder. Some years ago a
specimen measuring eighteen inches long was brought to
me dead, and on opening it I found it was packed with
seeds resembling those of the sycamore and another type

resembling pepper berries. I planted some of the former

and a few germinated and put forth the dicotyledons, but
died before its true leaves came, as the cold weather
set in. Most of the seeds and berries had been bitten,
I presumed that they must have fallen into the water
and then been taken as food. Most land tortoises are
vegetable feeders, and will hibernate if they have been
feeding well and are put into earth or moss.

These two groups are comparatively easily teat
with, they will feed readily and are not much subject to
diseases. Yet there are certain conditions to be aware of.
Crocodiles should not be fed entirely on meat, as there is
a tendency for their bones to become softened, no doubt
due to an insufficiency of lime salts. However, apart
from this, they are liable to injure their jaws and teeth in
biting at any hard substance introduced into the case,
and afterwards developing obstinate sores.. The Caspian
tortoise (Clemmys caspica) has often patches of diseased
bone in its carapace, and in separating, openings are left.

The Ophidia are very much more difficult to look

after, not so much on account of the heat or light as

EE —

EE ee EEE ee

REPTILE LIFE IN CAPTIVITY. 5

their susceptibility to diseases and the infectious nature
of these conditions. The most dreaded affection is a
membranous condition of the mouth which spreads to

‘

the fauces, and is known as ‘‘canker.’’ ‘This disease
gains entry to the mucous membrane inside the lips and
jaws through injury or abrasion. The animal becomes
ill-tempered and snappish, and ultimately dies, partly
as the result of starvation, through the inability of the
tissues to absorb food stuffs, and also partly from toxins
formed by the growth.

Another snake disease is known as “‘ casting disease,”’
and means that no sooner is the old slough cast than
another one begins, with the result that in a bad case the
unfortunate snake gets covered with a more or less thick
felted mass of partly shed scales. While snakes are
casting they will refuse food, and more especially when
in this chronic state. For canker I know of no actual
cure, but have always made a rule of isolating the
infected specimen and disinfecting the case thoroughly
to try and prevent the spread of this scourge. Casting
disease can at times be cured by bathing the animal in
water to which a little glycerine has been added.

Snakes are carnivorous reptiles and require water
for drinking and bathing. They show considerable
diversity in their methods of taking their prey, the
poisonous species strike and usually wait the victim’s
death. The constrictors kill their prey by encircling its
body with their own folds, while others, as the grass
snakes—Tropidonotus—swallow frogs alive, catching
them by a leg.

It is possible to get the pythons and boas to take
dead animals, but even then they will constrict them.
The capture and death of an animal by a constricting
snake is a remarkable process. Assuming that a rat is

6 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

put into a boa’s case, the snake glides towards it and
when quite close it will investigate it. Then, apparently
satisfied that all is right, it will retract its neck and
suddenly dart forward, usually catching the animal by
the ‘side of its head or neck, and a combined movement
follows, the snake draws its prey towards it and throws
a coil or two of its body round the luckless animal.
Should it struggle vigorously, additional coils are
brought into play, and it is held until dead, a matter
of two or three minutes. I do not think there can be
much, if any, pain as asphyxia sets in rapidly. It is
often said that a python pours saliva over its victim
before swallowing it, this I have never observed. The
animal when dead is released in order to be swallowed
head first, but no saliva is poured over it. Another
point often brought forward is the supposed fascination
and terror of the animals used for feeding these snakes.
There is no such thing. Whether the intended food is
rat, rabbit, or bird, no signs of fear are shown, but on
the contrary the victim will placidly feed in close
proximity to the snake. 7

The Pythoninae, as a group, are most interesting,
and become very docile in a short while. The common
boa (B. constrictor) is a frequently kept species, as it is
one of the handsomest and also one of the hardiest of
them all. It will readily take mice, birds, rats, or
rabbits, according to its size. It 1s very inquisitive
and will examine its surroundings or a stranger
minutely; it is somewhat nervous, but soon gets to know
its attendant. A rather remarkable fact in its economy
—at least in specimens up to six or eight feet—uis the
apparent absorption of all the lime taken in the bodies
of its victims. J mention this matter with some reserve,
although I have failed—spectroscopically—to detect hme

REPTILE LIFE IN CAPTIVITY. u

in the excreta. The excreta consist of hair masses,
entangled in which are a few small bones, which thus
have escaped the very active gastric juices, and almost
solid masses of uric acid.

A word of warning may be given to collectors and
others—and it is this: never put a freshly imported
specimen into a case with snakes which one knows are
healthy, always isolate the new arrival as it is quite
possible that it may have canker.

The black python (#. seba) of W. Africa is
frequently imported, and grows rapidly and does well in
captivity, as also does the Indian (P?. molurus). On the
other hand, P. regius is not a satisfactory feeder. I had
an unfortunate experience with a small snake of this
species once. It had not fed with me, and I had put a
rat weighing six ounces into the cage for a larger snake.
I may mention the small python also weighed six
ounces. It seized the rat and killed it, and with great
difficulty swallowed it, but only survived its meal some
twenty-four hours.

Snakes renew their epidermal shields periodically,
the young ones will cast every couple of weeks, and the
process becomes less frequent as the snake grows.

Many exaggerated stories are told of the large
animals taken by the constrictors as food, but it 1s
recognised that a snake can only swallow prey the
greatest diameter of which does not exceed that of the
snake’s neck by three times.

A boa in my possession increased from 25 lbs. to
7 lbs. in a year. Certain snakes will not feed readily in
captivity and must be fed artificially. This is best
done by means of a smoothly ground glass tube, which
is inserted into the snake’s mouth and chopped meat
gently pushed down. It is necessary to be very careful

8 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

during the operation as any laceration of the mucous
membrane may easily lead to canker. When a snake
bites it does not as a rule retain its hold, and this is true
for the constrictors, unless they have seized prey; and
this is directly the reverse of the next group, the lizards.
When the latter bite they usually hold on firmly,
grinding their teeth into one’s hand. It is advisable
to use great caution when the larger ones are being
handled. i

Lizards are rather more difficult to deal with in a
satisfactory manner in captivity.

As mentioned before, they require an amount of
sunshine before they will feed readily, quite indepen-
dently of the temperature of the case. The British
species, of course, do not require artificial heat, but
will feed well and remain active during the day, only
becoming dull at night time.

Most lizards are carnivorous and will eat worms and
insects of all sorts, the larger kinds will take mice or
small birds. Very few groups show such diversity of
form and special adaptability for various modes of life
as do these creatures. There are the water species,
mostly with compressed bodies and tails; the tree living
ones, with more or less long whip-lke tails and strong
claws; the flattened sand or desert species ; the curious
Geckos, with their palmate lamellated discs which enable
them to cling to the sides of walls and to run up to and
across ceilings. A remarkable peculiarity of many
lizards is the ease with which they part with a portion
of their tails, owing to a special mode of articulation,
and which no doubt aids them often in avoiding capture;
another use of the tail is as a balancing organ while
running or climbing.

Among the larger ones which are commonly

REPTILE LIFE IN CAPTIVITY. 9

imported we may mention the Monitors, these «re
usually long and lithe looking and somewhat
suggestive of the Ophidia. The head is flattened
and elongate, and armed with sharp teeth. A
curious power is possessed by this group, or to be
correct, especially noticeable in this group, and it is
that they seem to be able to bend their jaws when biting,
so that when one gets hold of a finger the anterior portion
of the jaw is not thrown out of action; and they hold
on most determinedly and give a very serious bite.
They will also fight amongst themselves. Their food
consists of meat, small mammals or birds.

The tuberculated Iguana is a very handsome species
of a bright leaf-green colour with darker marks which
are white edged. They feed best, in captivity, on
bananas, and require a branch to climb on. Like the
Monitors, they are not above using their long whip-like
tails as weapons of offence or defence.

The Mastigures are found in India and N. Africa,
and have compressed bodies which are covered with
small scales like grains of sand. Their tails are armed
with formidable spines.

As a rule, they are difficult to keep, but this summer
I kept several in a heated case in a small conservatory,
to which the sun had full access. The temperature used
to go up to 100° F.; and all fed ravenously on cabbage
and lettuce, and they drank water freely: Their
activity in this heat was wonderful.

The Geckos, being nocturnal, are seen best at night,
when they will run up the glass sides and dart about
after insects with great rapidity. The pretty little
Anoles have some power of changing their colour, but
not nearly as much as the Chameleons.

The stump-tailed Australian lizard (7rachysaurus

10 TRANSACTIONS LIVERPOOL RBIOLOGICAL SGCIETY.

rugosus) is twelve or eighteen inches long, and has large
scales like miniature fir cones on its head and back.
There are two poisonous lizards, known as Gila monsters
(Heloderma horridum and a second H. suspectum), they
certainly seem to have the power of killing small
mammals or birds with their salivary secretion.

In appearance the common Gila monster is a dusky
colour with broad yellowish bars across its back, and the
scales are tuberculated.

Many of these creatures become quite tame and
learn to know their attendant, and will watch for feeding
time. :
Tortoises present very little difficulties in the way
of keeping. Land species will usually feed readily on
green vegetables or fruit; the African genus Cinixys
has only eaten bananas with me. Needless to say, they
require an even heat. |

The water tortoises are among the most interesting
of this group; they are all carnivorous, with a very few
exceptions. |

The members of the American genus Chrysemys are
prettily marked when young, many of them having a
remarkable colour scheme, in which red, yellow, brown
or green predominates, and the general effect is very
striking. The best known is probably C. picta, which
has an olive carapace with yellowish or reddish stripes
outlining the shields of the back. Each marginal
shield has a circular red line on it, and the same colour
is seen on the under sides of these shields. The plastron
is yellow. They are to be had from the dealers, ranging
from one to three inches long, and will take small pieces
of meat or worms. Water living species have flat webbed
feet and swim well. |

A remarkable animal is the Alligator terrapin, a

REPTILE LIFE IN CAPTIVITY. ta

lumbering brute, with a large head, which it cannot
retract within its shell, a long scaly tail, and is very
vicious. A bite from a specimen a foot long would be
a very serious matter, as it has strong hooked jaws, and
takes every opportunity of snapping.

~The Trionyx group is remarkable in having a small
amount of bony shields in their carapaces and plastra,
and in being covered with soft skin. They are flattened,
and as the name implies, have only three nails on their
feet. Their nostrils end in a tubular prolongation,
and the horny jaws are sheathed in skin, yet the
larger members of this group are very savage.

A long experience with reptiles and amphibia las
afforded many opportunities of noticing remarkable facts
about them. On one occasion an agama, six inches long,
and a small stumpy-nosed crocodile shared the heated
ease with a large S. African bull frog (2ana adspersa),
The former was missed, and as suspicions rested on the
large frog—which was seven inches long from tip of
snout to vent—the amphibian was made to disgorge its
meal and the lizard revived. Ultimately, it and the
small crocodile were eaten by the huge frog, which
could also take mice as food. Most reptiles are
cannibalistic and will not hesitate to kill and eat their
own kind. A common chameleon caught and killed a
small anole, which was dashing about its case, before it
could be stopped. ,

Another chameleon caught a triton and bit it
severely; but in this case the biter was bit, as in spite
of being attended to at once the toxins of the newt
proving too much for the attacker; and the former died
within half an hour.

Chameleons are easily kept while the summer and
autumn months are in force, but as soon as winter

12 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

approaches, even if one has a stock of blue-bottles
hatched artificially, they usually die. It is necessary
to supply all reptiles with water, and certain species will
require spraying with water to keep them satisfactorily.
This is particularly the case with Mastigures.

Phrynosoma is not an easy species to keep, but very
much more difficult is the skink (Scincus officinalis),
which requires avery high temperature and an
abundance of sunlight. ,

One could expand notes and memories of strange
happenings while observing various species of ihe
reptilia while in captivity, but what strikes one most in
this group is the keenness of its members and their ever
alert watch on their surroundings, and they help to make
us realize what an earth peopled with the monsters of the
past must have been, and, incidentally, what small
chances the human race would have had in conflict with
them.

‘arog ‘eT ‘FJorgq Aq ydvaSoqoyd ve wor]
‘QSe] UWON OY} Wosy Woregg [OIsCToIg UG yOog oy, “T OLA .

THE
MARINE BIOLOGICAL STATION AT PORT ERIN

BEING THE
TWENTY-FIFTH ANNUAL REPORT

OF THE

LIVERPOOL MARINE BIOLOGY COMMITTEE.

Again we are happy in not having to record any
changes in the Committee or on the Staff. The year
has been a good one both in weather and in work, and
we are able to show an increased number both of
investigators in the Laboratory and of visitors to the
Aquarium. The new research wing added last winter,
and described fully in the last Report, has relieved
pressure, and has proved quite satisfactory and most
useful—especially during the Easter vacation. The
enlarged library is a pleasant room, and is useful, not
merely to accommodate the books, but as a sitting-room
common to all workers in the building for purposes
of reading, writing and occasional meetings. We have
now abundance of room for additional books on the
shelves; our present library of about 310 volumes and
620 Reports and pamphlets looks rather a meagre collec-
tion, and a considerable addition to the library of marine
biology is one of our most pressing needs.

Figure 1 (frontispiece) shows the Biological Station
in its present condition from a photograph taken last
Easter, after the additions were opened for work; and
fig. 2 gives the back view of the new wing, with its door
to the yard and the outside stair to the upper floor. The
library windows are seen in the latter figure under the
low-sloping roof, between the research wing and the
back of the aquarium. The ground plan (fig. 3) makes
the accommodation clear on both floors.

14 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The usual Haster Vacation Class in Marine Biology,
as a branch of Nature Study, was carried on with the
usual success during April, under the guidance of Mr.
Douglas Laurie and Dr. Dakin; and, in addition,
Professor Harvey Gibson held a course of lectures and
practical work on Marine Alge for Students of Botany.
Further details in regard to both these courses are given
below. Professor Cole again brought a contingent of
his senior students from University College, Reading.
We had a group of workers from Queen’s College,
Cork; and altogether six different Universities or
Colleges have been represented in the Laboratory.

Fic. 2. Back of the Biological Station, from the yard.

As on previous occasions, I shall first give the
statistics as to the occupation of the “‘ Tables’’ during
the year, then will follow the ‘‘ Curator’s Report,’’ and
the reports that have been sent to me by various

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MARINE BIOLOGICAL STATION AT PORT ERIN.

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16 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

investigators on the work they have done, and, finally,
I shall describe some of the researches upon which I
have been myself engaged.

Tue Station ReEcorp.
Sixty researchers and students have occupied the
Work-Tables in the Laboratories for varying periods
during the year, as follows :—

Dec. 27th to Jan. 9th. Professor Herdman.—Official.

March 24th to April 4th. Professor F. J. Cole.-—Educational. °
es Mr. H. L. Hawkins.—Echinodermata.
LF Mr. Malpas.—General. .

ae Miss Attride.—General.
March 24th to April 6th. Miss Davies.—General.

He Miss Freeman.—General.
March 27th to Aprit 10th. Mr. S. Mangham.—Nutrition in Marine Algz.
March 28th to April 11th. Mr. E. W. Shann.—General

i Dr. W. M. Tattersall—Embryology of Littorina.
xe Miss Kyffin.—General.

x Miss Stewart.—General.

ks Miss Payne.—General.

ae Miss Lindsey.—General.

a Miss Pearce.—General.
March 25th to April 29th. Professor Herdman.—Plankton.

Mr. W. Riddell.—Plankton and Polychaeta.
he Dr. Dakin.—Buccinum.

April 7th to 1th. Mr. R. D. Laurie.—Educational.
April 4th to 24th. Mr. H. G. Jackson.—Eupagurus.

- Mr. W. H. Evans.—Physiology of Invertebrates.

be Miss Latarche.—Biometry.

_ Miss Jackson —General.

rf Miss Jolley.—General.

oe Miss Scott.—General.

fs Miss Robinson.—General.

cs Miss Gleave.—General.

Bs Miss Coburn.—General.

a Miss Lewis.—General.

3 Miss Bamber.—General.

Lae Miss Robbins.—General.

Sh Miss Gill.—General.

Be Miss Firth.—General.

fe Mr. Daniel.—General.
April 7th to 22nd. Mr. R. H. Compton.—Marine Alge.
April 8th to 22nd. Miss Hood.—General.

Bs Miss Onions.—General.
April 13th to 29th. Prof. B. Moore.—Physiology of Invertebrates.
April 13th to 25th. Professor R. J. Harvey Gibson.—Educational.
April 13th to 29th. Mr. E. Whitley.—Physiology of Invertebrates.
April 14th to 25th. Mr. W. A. Gunn.—General.
April 15th to 22nd. Mr. J. C. Johnson.—Marine Alge.

huss; Miss Duke.—Marine Algze.

a Miss Dubbin.—Marine Alge.

a Mr. Mosley.—General.

. Mr. Megson.—General.

MARINE BIOLOGICAL STATION AT PORT ERIN.

April 14th to 24th.

June 3rd £0 6th.
July 7th.

July 10th to August 19th.
July 17th to 28th.

August 21st to Sept. 4th.
August 23rd to Sept. 15th.
August 23rd to Sept. 19th.
Sept. 5th to 18th.

Sept. 6th to 19th.

Sept. 7th to 19th.

Sept. 8th to 28th.

Sept. 23rd to 30th.

ies" Tables ’’

follows :—

Liverpool University Table :

Professor Herdman.
Dr. H. E. Roaf.
Dr. Dakin.

Mr. H. G. Jackson.

Professor R. J. Harvey Gibson.

Mr. Laurie.

Miss Knight.—Marine Alge.
Miss Howlett.—Marine Alge.
Miss Edge.—Marine Alge.

Miss Stubbs.—Marine Algz.
Miss Galloway.—Marine Alge.
Miss Grundy.—Marine Alge.
Miss Gleave.—Marine Alge.
Miss Molyneux.—Marine Alge.
Miss Beardsworth.—Marine Alge.
Professor Herdman.—Ofificial.
Professor Herdman.—Plankton.
Mr. W. Riddel!.—Plankton.

Dr. H. E. Roaf.—Physiology of Invertebrates.

Mr. R. A. Wardle.—General.
Mr. Holden.—Marine Algz.
Mr. J. C. Waller.—General.

Professor B. Moore.—Bio-Chemistry of Echinus.

Professor Herdman.—Plankton.
Mr. W. A. Gunn.—General.
Mr. Bury.—General.

Mr. W. Riddell.—Plankton.
Dr. Dakin.—Buccinum.

Mr. E. Hamilton.—General.

in the Laboratory were occupied as

Mr. E. Hamilton.
Mr. Gunn.
Professor B. Moore.
Mr. Whitley.

Miss Latarche.

Mr. W. H. Evans.

Liverpool Marine Biology Committee Table :—

Mr. 8S. Mangham.
Mr. Mosley.

Mr. Riddell.

Mr. R. H. Compton.

Mr. Megson.
Mr. Bury.
Mr. J. C. Waller.

Manchester University Table :—

Dr. W. M. Tattersall.
Mr. Holden.

Miss Payne.

Mr. E. W. Shann.
Miss Kyffin.

Miss Lindsey.

Mr. R. A. Wardle.
Miss Stewart.
Miss Pearce.

Birmingham University Table :—

Miss Hood.

Miss Onions.

University College, Reading, Table :—

Professor F. J. Cole.
Miss Attride.
Mr. H. L. Hawkins.

Miss Davies.
Mr. Malpas.

Miss Freeman.

The following senior students of Liverpool Univer-

sity,

and of Queen’s

College, Cork,

occupied

the

Laboratory for periods varying from a fortnight to three

weeks during the Easter vacation, and worked together

18 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

under tae supervision of Professor Harvey Gibson, Dr.

Dakin and Mr. Laurie.

Miss Jackson. Miss Jolley. Miss Scott.

Miss Robinson. Miss Gleave. Miss Coburn.

Miss Lewis. Miss Bamber. Miss Robbins.
Miss Gill. Miss Firth. Mr. Daniel.

Miss Knight. Miss Howlett. Miss Edge.

Miss Stubbs. Miss Galloway. Miss Grundy.
Miss F Gleave. Miss Molyneux. Miss Beardsworth.
*Mr. J. ©. Johnson. *Miss Duke. *Miss Dubbin.

In addition to these workers, the Station has been
visited during the year by several travellers interested
in Natural Science and Education, by various groups
of more senior school pupils with their teachers, and by
members of scientific societies.

We have been endeavouring for some years to bring
the Aquarium and Museum more closely into connection
with the educational system of the Island, and I am
glad to say that a beginning has now been made. Mr.
Ashton, the Headmaster of the Secondary School, sends
eighteen of his boys every Wednesday afternoon for a
lecture and practical demonstration, from our Curator,
lasting from 2-30 to 4 o’clock. Mr. Chadwick writes to
me after the first of these: —‘‘ I began with a lesson on
the Protozoa, and after a short lecture, with plenty of
black-board sketches, I got out a couple of microscopes
and showed the boys an Amceba and a number of
Foraminifera and Radiolaria. They all bring notebooks,
and Mr. Ashton requires them to write out, afterwards,
an account of each lesson. Next week I shall take
animal cells, so that they may understand something
of the structure of the various animal tissues. Then I
shall take each group of Invertebrates in ascending
order.”’

This ought to do good to the boys and help the
school. It is a small beginning in that general intro-

*From Queen’s College, Cork.

MARINE BIOLOGICAL STATION AT PORT ERIN. 19

duction of the methods and results of science to the
rising generation which is so desirable in the interests of
national progress.

Curator’s Report.

Mr. Chadwick reports to me as follows :—

‘““T again have the pleasure of recording an increase
in the number of researchers and students who have
resorted to our laboratories, the total number this year
being sixty. The Universities and other institutions
represented were almost the same as those of last year,
with the addition of Queen’s College, Cork, and the
Technical School, Huddersfield. Of the sixty workers,
forty-two occupied tables at the one time, during the
Easter vacation; and, in spite of the increased
accommodation afforded by the new research laboratory,
it was found necessary to utilise the museum gallery in
order to avoid overcrowding and its attendant incon-
veniences.

“The weather during the Spring vacation was, on
the whole, favourable for out-door work, and much shore-
collecting was done under the guidance of Professor
Herdman, Professor Harvey Gibson, Professor Cole, Dr.
Dakin and Mr. W. A. Gunn. Several visits were paid,
by boat from Port St. Mary, to the caves-in the
neighbourhood of the Sugar Loaf Rock, and were
greatly appreciated by the senior students who
participated in the exceptional opportunities of collecting
afforded by these expeditions. In addition to the
practical instruction given in the laboratories during
the day, evening lectures were given by Dr. Dakin, on
‘Plankton,’ and by the Curator, who gave an account
of his studies on the local Echinoderm larvae, illustrated
by a large series of lantern slides photographed from his
own original drawings.

20 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

‘‘The various branches of the work of the Station,
including the collection and preservation twice a week of
Plankton gatherings in the bay, have been carried on
successfully throughout the year. The collections have
been forwarded periodically to Mr. Andrew Scott to
be examined in connection with Professor Herdman’s
‘Intensive Study of the Plankton.’

‘About a year ago Mr. G. H. Wailes, who is com-
pleting the Monograph on the British Freshwater
Rhizopoda begun by the late Mr. James Cash and
published by the Ray Society, asked me to send him
samples of damp moss from water courses and boggy
places in the neighbourhood of Port Erin. I did this,
and Mr. Wailes has kindly supplied the subjoined list
of Rhizopoda and Desmidiacee found in a quantity of
moss from the little cascade which falls over the cliff
close to the old harbour workshops. Very little is known
of the freshwater Rhizopoda of the Isle of Man, and the
list of species found by Mr. Wailes seems to indicate a
promising field of research.

RHIZOPODA.
Lososa. FILosa.

Arcella vulgaris. Euglypha alveolata.

BS » var. gtbbosa. sa laevis.

» discordes. a » » with apical spines.
Centropyxis aculeata. Sp. nov.

arcelloides. Trinema enchelys.

Difflugia oblonga. lineare.

99 39

es globulus.

aN constricta.

ae pristis.

~ lucida.

rubescens.
Pontigulasia compressa.
5 bryophila.

Pyxidicula operculata.
Quadrula symmetrica.

= rregularis.
Heleopera petricola.

Cosmarium speciosum.
be formulosum.

var. lacustris.

Cyphoderia ampulla.

>» 5 var. major.

» cp other varieties,
Sphenoderia dentata.
Pamphagus hyalinus.

DESMIDIACE 4A.
Cosmarium laeve.
= Botrutis.

MARINE BIOLOGICAL STATION AT PORT ERIN. 21

““The number of visitors to the Aquarium during
the year—13,200—shows a slight but gratifying increase
compared with last year. No exceptionally large
attendances on single days were recorded, but the daily
average attendance improved substantially towards the
end of the season. The number of copies of the ‘ Guide
to the Aquarium’ sold—over 700—again shows a con-
siderable increase, and bears testimony to the well-
sustained interest of the more intelligent visitors. A
new edition will be required during the coming year, and
is now in preparation.

‘** Visits have been paid to the Station by the Douglas
Progressive Debating Society and the Douglas Sunday
School Teachers’ Association, in addition to seven visits
of parties of boys and girls from local and other insular
elementary schools. The Curator was present on all
occasions, and generally gave a short illustrated lecture
in addition to an explanation of the contents of the tanks.
The Station was visited in July last by MM. Docteur
Armand Geoffrey, Médecin de la Marine, and Robert
Cayrol, Enseigne de Vaisseau, Boulogne-sur-Mer; and
in September by Professor HK. W. MacBride, F.R.S. On
September 14th a meeting of the Isle of Man Natural
History and Antiquarian Society took place at the
Biological Station. The chair was occupied by the
Deemster Callow, Chairman of the Fishery Board, and
an address was given by Professor Herdman on ‘ Science
in relation to the prosperity of nations, and the need of
a more wide-spread education in the methods of science.’

‘Owing to the comparatively small number of adult
plaice in the spawning pond during the hatching season,
the number of fry set free in the neighbouring seas was
considerably less than that of last year. Favoured by
the comparative absence of boisterous winds, the pond

22, TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

was thoroughly skimmed by the Assistant Curator on
forty-five of the fifty-eight days over which the season
lasted. The largest number of eggs obtained on one day
was 306,000 on April 15th, but collections of over
300,000 were made on three previous days. The total
number for the season was 7,462,000, and the total
number of fry set free in the sea was 4,929,600. The
majority of the young fish were liberated by Professor
Herdman from his yacht ‘ Runa,’ at a distance of about
five miles from land. |

‘‘ Figures 4 to 9 show aseries of stages in the develop-
ment of the young plaice from the newly hatched
larva (4) through the post-larval metamorphosis (5 to 8)
to the young flat-fish (9). They are from photographs
taken at Port Erin by Dr. Francis Ward.

‘“The numbers of plaice eggs collected and of larvae
set free during the past season were as follows :—

Eggs collected. Date. Larvee set free. Date.
121,800. March 2 90,300 March 21
135,400 ie 3 166,400 sis ghee

59,800 4 4 47,200 Be ae
108,000 7 6 85,000 a
187,000 a | 135,500 ene)
116,500 Min 8 91,300 » 30
169,000 2) LALO 135,500 5. 0
125,000 see abl 101,000 ie 2108
191,000 Ss le 164,700 April 4
192,250 cies Le 164,000 B: 4

99,700 ‘cipal 79,700 5 7
174,300 er Ks) 139,700 os |
159,600 LS 117,000 Ss
199,500 + 8120 168,000 Shei
176,400 aay all 139,700 Were
203,800 ire 48s 162,700 sth wal
175,300 ae 140,700 ‘nen be
198,500 5. 24 147,000 5 ile
135,500 ast PZD 90,000... re |)
202,600 te 7 loGA00 ees ¢ JS
188,000 ae ae 137,200 on
312,900 3. ead 241,300 18

MARINE BIOLOGICAL STATION AT PORT ERIN.

Eggs collected. Date. Larve set free. Date.
347,000 March 31 220,000 April 22
143,800 Jsjayet bigaal | 94,500 A
315,000 ae 3 220,000 hat ee
235,200 is 4 160,000 beky 0
219,000 sd 5 137,000 Bese
179,500 a 6 112,000 Sue
195,300 i 7 136,500 a ae
195,000 ie 8 124,000 ile
216,300 re AG 132,000 May I
197,400 sear At 96,000 +5
174,300 ea wiki get 90,000 iy 1
140,700 5 eal bo 72,500 i 1
356,200 pe NING 150,500... Pe 1
233,000 a ARG, 89,500. ... - 6
129,000 ite ke. 50,000... Ep 6
137,500... ee 19 GE OOO on 3, 5 6
(0 oy eee At) 20,000... - 6
141,700 ... + ey DE TOO eae 3 6
84,000... a 29,400... te
C2000". +... 5) tC 24,300... yt oe
T6800: ;. ee. “Di
ae. (st spain eae ar
7,462,000 4,929,600

‘* When the plaice hatching season was over (fig. 10),
the Curator turned his attention to lobster culture, and
by great and persistent effort over twenty ‘ berried’
female lobsters with eggs in various stages of ripeness
were acquired from local fishermen. The eggs of one of
these began to hatch out on June 21st, and experiments
in rearing were at once begun, while some of. the larvae
in the first stage were set free at points where it was
thought that suitable ‘cover’ would be found.
Altogether 7,450 larvae were hatched, and every effort
was made, by experiment on old and new lines, to rear
them through their early stages, but without tangible
success. All the difficulties of past years were
experienced, and it is now abundantly clear that the

i
'

TRANSACTIONS

LIVERPOOL BIOLOGICAL

SOCIETY.

PSETEE eae

i = ries

MARINE BIOLOGICAL STATION AT PORT ERIN. 25

propensity of the berried lobsters to shed their ripening
eggs cannot be obviated when they are confined in small
tanks. The same conditions appear to apply to the
health of the larvae. Of 1,400 larvae placed in the large
concrete tank in the hatchery only six reached the
lobsterling stage; while a considerable percentage of
900 placed in the spawning pond were seen
swimming at or near the surface until the third (and in
a few cases the fourth) larval stage was reached. Many
of these would probably have reached the lobsterling

Fic. 10.—The Hatching Tanks when used for rearing lobsters
in the Summer.

stage but for the presence of the fish in the pond. It is,
no doubt, probable also that the abnormally high
temperature to which the water in the pond was raised
on many days in July and August by the exceptional
heat of the sun this summer had a prejudicial effect on
the lobster larvae. It is a curious fact that all our
experiments in lobster rearing in this and past years
have afforded striking illustrations of the ‘ survival of

26 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the fittest.’ - Whenever some hundreds of larvae are
confined together in the comparatively small space of a
tank, about 2 per cent. or 3 per cent. only reach the
lobsterling stage. It apparently matters not whether
they are artificially fed or left to exercise their canni-
balistic propensities—the result is in all cases practically

Fic. 11. Newly-hatched young lobster. From a photograph by
Mr. Edwin Thompson.

the same. No difference in the rate of mortality at the
periods of larval ecdysis has been observed between
artificially fed larvae and those which have lived entirely
upon their weaker brethren. The mortality at the
periods of ecdysis was exceedingly heavy amongst the
1,400 larvae in the hatchery tank, though they were fed |
at frequent intervals and supplied with abundance of

circulating water. a4

MARINE BIOLOGICAL STATION AT PORT ERIN. 27

OTHER REPORTS ON WORK.

Professor Harvey Gibson writes as follows :—

“During the Easter vacation a course on Marine
Algz was given at the Station, attended by over twenty
advanced students from the Botanical Department of
the University, by two students from the University of
Birmingham, three from University College, Cork, and
one from the University of Cambridge. The course con-
sisted in systematic collecting in Port Erin Bay, Port
St. Mary, and elsewhere. The plants collected were
examined in the laboratory during flood tide, and each
evening a short lecture was given on the morphology and
life-history of representative forms. In addition to the
identification of species already recorded. from the
district, several new forms were found and material was
gathered for subsequent research. It is hoped at an
early date to publish in the ‘Transactions of the
Biological Society ’ some notes upon these new species,
and on certain morphological points which have not
hitherto been described. One hundred and fifty-seven
species in all were collected, among which the following
are the more important novelties : —

CHLOROPHYCEAE. PHAEOPHYCEAE.
Prasiola stipitata. Ectocarpus granulosus,
Enteromorpha percursa. 5 hincksit.
Endoderma witrockii. Cladostephus verticillatus.
Epicladia flustrae. Ascocyclus leclancheri.
Cladophora nuda. Leathesia difformis.
Bryopsis hypnoides. Sporochnus pedunculatus.

Fucus ceranoides.
RHODOPHYCEAE.

Bangia fuscopurpurea. Dilsea edulis,
Helminthocladia purpurea. Petrocelis cruenta,
Callophyllis laciniata. Peyssonnelia dubyi.
Catenella opuntia. Hildenbrandtia rosea.
Sphaerococcus coronopifolius. Rhodomela lycopodioides.
Champia parvula. Laurentia obtusa.
Delesseria hypoglossum. Polysiphonia violacea.
Ceramium gracillimum. 7 byssoides.

9 acanthonotum Nemalion multifidum.

28 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

‘‘Some of the above were collected by Miss M.
Knight, B.Sc., and Miss H. Coburn, B.Sc., at Peel
during the Summer vacation.’’

During July and August Dr. H. E. Roaf investi-
gated the effect of the concentration of oxygen and free
carbon dioxide on the rhythmical movements of marine
organisms. He writes as follows :—‘‘ The object was to
find out if these movements, like those of mammalian
respiration, were mainly influenced by the partial
pressure of carbon dioxide and not appreciably affected
by the concentration of oxygen. In the event of carbon
dioxide exercising the main influence, it was further
desirable to discover if this effect were due to an increase
of acidity or to the increased partial pressure of carbon
dioxide. The experiments were carried out by measuring
the rate of the rhythmical movements of rock barnacles
(Balanus balanoides) and the rate of the gill movements
of a small fish (Cottus scorpius) under different
conditions. The results for barnacles suggest that the
lower the oxygen concentration the slower are the move-
ments, and with carbon dioxide increasing concentration
makes the movements more sluggish. With the fish the
reverse holds, namely, that the low concentrations of
oxygen and high concentrations of carbon dioxide both
increase the rate of movement. The influence of oxygen,
however, is so slight that it approaches the limit of
experimental error. In both cases the action of carbon
dioxide was due to the increase of acidity (hydrogen ion)
with increased concentration of carbon dioxide. It is
hoped to publish full details of this investigation and its
results in a short time.”

Dr. Tattersall, of Manchester University, reports : —

‘‘ During the Kaster vacation I spent a fortnight at
the Port Erin Biological Station, in a further attempt to

MARINE BIOLOGICAL STATION AT PORT ERIN. 29

rear the eggs of the periwinkle (Littorina littorea)
through the various stages of their development. Very
little success attended my efforts, and I made no progress
with the work. I hope to renew the attempt next
spring. I am able to add one species of Crustacea to
the Port Erin list and to the L.M.B.C. district. On
April 11th last, four specimens of Schistomysis arenosa,
G. O. Sars, were taken in a small hand dredge dragged
lightly over the surface of the sand in the centre of the
bay. This species is known from the Clyde area, from
several places on the Devonshire coast, and from
Blacksod Bay in the West of Ireland. As its name
implies, the species normally lives buried in fine sand.”’

Professor Benjamin Moore and Mr. Edward
Whitley were engaged during the Easter vacation upon
an enquiry into the bio-chemistry of the reproductive
organs of Hcehinus, which was continued by Professor
Moore during the summer months, and a large amount
of material was accumulated and extracted in addition
to the work actually carried through at the Port Erin
Station. This material is now being utilised in the
Bio-Chemical Laboratory at the University of Liverpool
for the study of two distinct problems in metabolism by
Mr. N. G. 8. Coppin, B.Sc., and by Mr. Alfred Adams,
M.B., Ch.B.

Professor Moore reports : —

“At the outset the purpose of the research was to
discover whether the reproductive organs of Hchinus
esculentus contained any of those more simple represent-
atives of the protein classes, termed protamines and
histones, found by Miescher, Kossell and others in the
sperm of fishes, and by Matthews in another form of
echinoderm. A body closely resembling the arbacin of
Matthews was isolated and its properties studied both at
Port Erin and Jater at Liverpool.

30 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

“The primary object in obtaining such bodies which
possess strongly alkaline or basic properties, was to test
whether they acted as stimulants to cell-division,
following up the previous work at the Port Erin
laboratory of Moore, Roaf and Whitley a few years ago.
In this respect we were disappointed, the histone isolated
being found to have no exciting or accelerating action on
the speed of division in the fertilised Hchinus egg. But,
at the same time, the physico-chemical properties of this
body have been found very interesting. The osmotic
properties and pressures developed by its solutions have
been studied, and it has been shown to pass into true
solution with a high osmotic pressure, demonstrating
that these simpler proteins really do possess a _ less
complex molecule.

‘“While this work was in progress it was observed
that the gonads, both male and female, were very rich
in fats, which were incidentally removed as a
preliminary to obtaining the protamines and _ histones.
Also, it was noticed that a batch of gonads obtained
from Echini which had been kept for about a week
without added food in the tanks, apparently yielded less
oil on extraction than the gonads of specimens taken
fresh from the sea.

‘“Vhis suggested the idea that the gonads, in
addition to their more obvious function, might also act as
metabolic organs, like the hepato-pancreas of molluscs
and arthropods, or the liver of mammals. It was,
therefore, determined to study the chemistry of the
gonads at different seasons of the year, and especially to
estimate the nature and amount of metabolic products,
such as fats and glycogen present at different times.

‘With this object in view batches of Echini were
assembled in August and September. Some lots were

MARINE BIOLOGICAL STATION AT PORT ERIN. a

killed and the gonads extracted in the fresh condition,
others were kept fed in the tanks and the gonads
extracted, and still others were kept for a period without
food and then killed and the gonads extracted.

‘*So much can already be said that, so far from
representing an exhausted gland at this time of year,
the gonads in both sexes are as large as in the breeding
season and contain both fats and glycogen in large
amounts. The glycogen content is being investigated
by Mr. Coppin, while Mr. Adams is determining the
nature and amount of the fats.

“One important point to be determined is whether
the amount of metabolic material stored up varies with
the state of nutrition, and whether such variations occur
apart from the breeding season.”’

Mr. W. Riddell, M.A., was at work at Port Erin
during both the Easter and the summer vacations,
assisting me with the Plankton investigation; but in the
intervals of that work he took up one or two other
matters. He continued his systematic work on the
Polychaeta of the neighbourhood, and to the list given in
last report he has been able to add Nereis virens, Glycera
siphonostoma and Notophyllum foliosum, the latter
dredged off Dalby in September, 1911. Mr. Riddell
also commenced an investigation into the nature of the
disease which has been causing much damage to the
plaice in the spawning pond during the last few years,
but this work has hardly been carried far enough as yet
to warrant any definite statement.

Dr. W. J. Dakin reports to me as follows on his
research work : —
“During the Easter vacation I conti
uring the HKaster vacation [ continued some of
the osmotic pressure experiments commenced at
Helgoland. These experiments had shown that the

32, TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

blood of teleosts from the aquarium tanks had a lower
osmotic pressure than blood from freshly-caught (living)
fish from the sea. There was a possibility that this
difference might be due, in an indirect manner, to
differences in hydrostatic pressure. In order to
determine the factor bringing about the abnormal con-
stitution of the aquarium fish, a number of living plaice
were taken out to sea in the ‘ Runa,’ and half were
lowered in a perforated box to the bottom in about 20
fathoms of water, whilst the remainder were kept at the
surface in a floating box. Full details of the methods
employed will appear in the paper (to be published in
the ‘Internationale Revue d. Hydrobiologie’). The
osmotic pressure of the blood from both the batches of
fish was found to be identical. Evidently, therefore,
differences in hydrostatic pressure are not responsible
for the differences previously observed in the blood, and
other aquarium conditions must be examined in order to
find the cause of the abnormalities.

‘“In connection with my work on the Whelk
(Buccinum undatum), which was continued at Port Erin
both in the Haster vacation and during September, some
details may be of interest. It has already been shown
that one can almost always be certain of finding stages
of an interesting Coccidian parasite in the renal organ,
and that for class work this may be regarded as a good
example of a Sporozoon easily obtained. Another
parasite occurs in the stomach and rectum—namely, an
endo-parasitic Turbellarian (Grafilla buccinicola). As
almost every whelk appears to be infected and contains
on an average about a dozen specimens, this may also
be taken as a convenient type for laboratory teaching
purposes. It is particularly valuable, since endo-
parasitic Turbellaria are by no means common. The

MARINE BIOLOGICAL STATION AT PORT ERIN. 33

parasite was discovered in Port Erin whelks fourteen
years ago, by Jameson, and it does not seem to have been
recorded elsewhere. It is interesting to find it turning
up still in the same place. Jameson states that it occurs
in the kidney and kidney duct, but those that I have
found were always in the stomach and rectum. As the
kidney opens directly to the exterior I cannot quite
understand what Jameson meant by kidney duct.

‘““ Most of the detailed work on the anatomy of the
whelk has now been completed, and I hope to have the
MS. ready for publication in January.”’

During part of the Easter vacation Miss M.
Latarche, B.Sc., made a _ preliminary _ bio-metrical
investigation of the variation in the shells of the common
limpet (Patella vulgata). These were taken from three
different localities, viz.:—Port Erin, Fleshwick Bay
and Port St. Mary.

Comparisons were made between sets of shells
taken—

(1) From high and low water marks;

(2) From Carboniferous limestone and Cambrian

slate ;

(3) From more exposed and more sheltered waters.

The variation occurred chiefly in the height of the
cone compared with the breadth. ‘The shells of those
taken from about low-water mark were found to be much
flatter than those taken from near high-water mark.

The nature of the rock on which the Patellae lived
did not affect the height of the shells. Those found on
the limestone at Port St. Mary were more markedly
ridged and were of a lighter colour. Intermediate links
between these and the darker, smoother, shells of Port
Erin were found to exist in both localities.

34 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Contrary to expectation there was no difference in
height between shells taken from the more exposed and
the more sheltered places, if taken at the same water
level. The two sides of the Port Erin breakwater served
as splendid collecting grounds for material for this
purpose.

Mr. H. G. Jackson, B.Sc., in addition to giving
me some assistance in taking the observations and
plankton gatherings at sea during the Easter vacation,
started an investigation of the complete anatomy and
histology of Pagurus bernhardus, the Hermit Crab. Mr.
Jackson is now continuing that work during the winter
in the Liverpool Laboratory, and the results will be
published, when completed, as an L.M.B.C. Memoir.

Professor F. J. Cole, with two members of the Staff,
Mr. H. L. Hawkins and Mr. A. H. Malpas, and three
of his senior students from University College, Reading,
worked at Port Erin during the Easter vacation.
Professor Cole writes to me, as follows, in regard to their
work : — | :

“The visit to Port Erin, last Easter, of our College
party of six was partly for educational and partly for
collecting purposes. The College has now provided for
the Easter class in the estimates, so that it will now
become an annual event. Its importance and popularity
with the students are too manifest to be dwelt upon, and
we expect to have a larger party working at the Station
next Easter. We were able this year to add two species
to the local fauna—Nereis virens, from Port Erin Bay,
and a peculiar Rhizocephalon, related to Peltogaster,
but still undetermined, which was found parasitic on a
large Galathea. We brought back a quantity of material
for the Museum, including a number of successful
injections of Echinoderms, Hledone and Fishes.”

MARINE BIOLOGICAL STATION AT PORT ERIN. 35

PLANKTON INVESTIGATION.

These observations have been carried on in much
the same way as in the previous four years, with the
kind help of various Assistants; and the new yacht
(S.Y. “‘Runa,”’ 95 tons, see fig. 12) has proved most
satisfactory, and very comfortable for work at sea.

EE —————— LL =

Fic. 12. Prof. Herdman’s fishing-yacht ‘‘ Runa,” from a photograph taken by
Dr. C. Macalister, at Shetland, on August 10th, 1911.

During the Easter and the Summer vacations Mr.
Riddell and Mr. H. G. Jackson helped me in the
observations on board the yacht, Mr. Chadwick did some
of the work of preserving the catches on shore and also
supervised the collections made from the bay during the
remainder of the year, and Mr. Andrew Scott is now
examining all the gatherings in detail with the
microscope.

Altogether 119 samples were collected at sea from the
yacht in the EHaster vacation, and 84 in the latter part
of August and in September [in the earlier part of the

36 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

summer vacation I was engaged in taking observations
further North, off the West Coast of Scotland]; while
about 150 gatherings were obtained from the bay during
the remainder of the year.

The year has been a somewhat unusual one in
weather, and that may have had some effect upon the
plankton. The material has not yet all been examined,
and it is still too early to make any very definite
statement, but the following provisional remarks, giving
the impressions formed at the time of collecting, may
be of some interest :—

In April, at Port LEHrin, Biddulphia and
Coscinodiscus were well represented, and also Chaetoceras
decipiens and C. teres.

At Piel, in the Barrow Channel, Mr. Scott tells
me, between April 15th and 19th, the plankton was very
rich in Chaetoceras and Rhizosolena, and a few
Thalassiosira nordenskioldi were noticed early in March,
but this and some other typical spring Diatoms, such as
Lauderia did not occur in any quantity at Port Erin
this Easter. Noctiluca was still living in the sea at Port
Erin up to the end of January, an unusual circumstance
due, perhaps, to the mild winter.

May 13th.—The vernal Diatoms now appeared in
quantities at Port Erin (calm weather with a marked rise
in temperature).

May 16th.—Tow-net gatherings large, and consisted
almost entirely of Diatoms (weather continues calm and
the increase in temperature is maintained).

May 19th.—Diatoms occurred in very large
quantities, especially in the fine net.

May 22nd.—Catches rather smaller, but Diatoms
still in abundance, even in the vertical net (weather still

fine and warm).

MARINE BIOLOGICAL STATION AT-PORT ERIN. 37

May 25th.—Diatoms much less numerous. Fine net
had only about one-tenth, or less, of the gatherings on
19th (no obvious change in weather conditions).

All the above large catches of Diatoms consisted
almost entirely of Chaetoceras. It was not until a week

later that Rhizosolenia (fig. 13) made its appearance. It
reached its maximum early in June, and then gradually
died off. By the beginning of July the Diatoms had

practically disappeared.

Fig. 13. Diatom Plankton, consisting Fria. 14. Copepod Plankton, consisting
mainly of Rhizosolenia semispina. wholly of Calanus helgolandicus.

I insert here a short list just received from Mr.
A. Scott, giving the quantity of plankton and _ his
estimate of the total number of Diatoms present in each

38 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

haul of the fine net, taken during the month of the
Diatom maximum. |

Date. Quantity in c.c. ~ Total Diatoms.
May 1 ": 2-5 Bis 43,360
UD Tea nN 1-0 i) 10,610
sp uO sae 6°5 sid 525,68C
he 5 gt oe 30:2 te 19,118,000
5 AG Re 60:2 im 54,141,500
eG it 54:5 cm 34,447,500
Scum ais 30°5 ae 27,775,000
see 25 see 8:3 a: 2,504,500
5 00 a 14:8 fee 22,023,100
June 1 a 11:3 ae 4,620,000
Rees an 24-7 ec 13,182,000
55 5 wwe 12% dis 2,143,000

On July 5th Port Erin Bay was invaded by an
exceptional swarm of the Copepod Calanus helgolandicus
(fig. 14), such as for some years now has always appeared
for a few days about this part of the summer. In 1909
there was an enormous Calanus swarm on July 17th
to 19th.

After being absent or rare during July, August and
most of September Diatoms made their appearance
again, for the Autumn visitation, towards the end of
September, and were very abundant in all the
gatherings during the first week of October. They
continued to be present in quantity during October, the
high numbers keeping up longer than usual. Both species
of Biddulphia (B. mobiliensis and B. sinensis, see fig. 15)
occurred in quantity in October, and occasionally in —
September, an unusually early appearance for B. sinensis, —
which seems to be in very vigorous condition this year.
Readers may be reminded that this is the species from the
Far East, which made its appearance in European seas
eight years ago, and is rapidly spreading along our

MARINE BIOLOGICAL STATION AT PORT ERIN. 39

coasts. Fig. 16 shows the two species of Biddulphia under
a higher magnification.

Our B. mobiliensis (Fig. 16, a) approaches the form
“regia”? regarded as a distinct though allied species by

Fie. 15. Plankton showing Biddulphia sinensis and B. mobiliensis.

Ostenfeld. B. sinensis (Fig. 16, b) seems to be of more
elongated form in our district than in Ostenfeld’s figures.

The autumnal Diatoms finally disappeared at the
end of October, and since then the plankton has remained
relatively small in quantity. A detailed account of the
plankton catches of the year will, as usual, be given by
Mr. Andrew Scott and myself in the Lancashire
Sea-Fisheries Laboratory Report, early in 1912. We
may remark here, however, that the figures given above,
for May, are unusually large, and that the increase from

40 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the ten thousand on May 4th to over fifty-four millions
on May 16th, is most rapid. The most abundant species |
were Chaetoceras debile and Ch. sociale. On May 16th

Fie. 16. Biddulphia mobiliensis (a) and B. sinensis (6).

the first of these species reached thirty millions and the
second 12 millions in the standard fifteen minutes’ haul
with a small net one foot in diameter of mouth.

ExtenDED INVESTIGATION Up THE West Coast.

In last year’s report I pointed out that our
‘plankton ’’ of
the western coasts of the British Islands was very

knowledge of the minute floating life or

incomplete because of a great gap extending from the
North of Scotland down to the Irish Sea—a gap which
neither the International Observations, on the one hand,
nor those of the Scottish and the Irish Authorities, on
the other, seem to fill up.

‘With a view to obtaining some data that may, in
part at least, bridge this gap, and possibly throw light

:
,
,

MARINE BIOLOGICAL STATION AT PORT ERIN. 4]

upon the causes of the seasonal changes in the plankton
of the Irish Sea, for several years back, during the

‘summer vacation, I have taken plankton hauls, both

vertical and horizontal, from the yacht at many
localities amongst the islands and lochs of the West of
Scotland, as far north as Portree in Skye, and as far out
to the west as the island of Barra. I was able to show

Fic. 18. Phytoplankton.

Fie. 17. Zooplankton.

in the last report, from these Scottish gatherings, that
the state of affairs at that time of year is somewhat
different from that in the Irish Sea. At some spots in
Hebridean Seas, for example, very large Phyto-plankton
hauls may be taken year after year in July—at a time
when in Manx waters the hauls are for the most part
comparatively small, and are all composed of Zoo-

plankton (figs. 17 and 18).

42 TRANSACTIONS LIVERPOOL BICLOGICAL SOCIETY.

During this last summer (July and August, 1911) I
devoted a longer time than usual to a more detailed
survey, with both bottom and surface nets, of a
considerable area off the West and North of Scotland.
The larger yacht we now have available for such work
renders it possible to go further, take a larger party and
remain out longer. Mr. Wm. Riddell acted as my
Assistant on part of the cruise, and will help in working
up the details of the material collected for a later report.
I can only at present give a brief preliminary account
of the results obtained. Our observations extend from
the Irish Sea as far North as Noup of Noss in Shetland
(from 54° N. lat. to 60° N. lat.) and as far West as
Castle Bay in Barra. They include 152 observations of
the sea-temperature, and 142 of the salinity. On August
22nd, when crossing from the south end of Cantyre to
the North of Ireland, a series of nine temperature and
salinity observations were taken, one every hour during
the most important part of the traverse; and on the

following day when crossing from Larne in Ireland to

Port Erin, another series of eleven hourly observations
was taken. During these two months (July 7th to
August 23rd) the temperatures varied from 11°2°C. to
17°8° C., and salinities from 1018 (= 22°69 °/,.) to 10276
' (= 84°87 °,,), the latter reading being a very high

salinity for British seas. It was recorded on August —

12th in the open sea to the Hast of the Shetlands, but
nearly as high a reading was obtained off Fair Island,
off North Ronaldsay and elsewhere in the Orkney seas,
and 1:027 was obtained on July 18th and 14th, off
Canna and Rum on the West of Scotland.

It would be premature, until the samples have been
more fully investigated, to make any positive statements
as to how this year’s observations compare with those

OO ee ee ee eee ee ee connie

MARINE BIOLOGICAL STATION AT PORT ERIN. 43

of previous summers; but this much may be said that the
impression produced at the time of collecting (fig. 19)
was that during July and August the Diatoms were less
in evidence, that the Phyto-plankton was less in amount

Fia.19. Taking Plankton and Salinity observations on the ‘* Runa,”

and less widely extended, and that a larger proportion
of the present collection is Zoo-planktonic.

In addition to the investigation of the plankton a
certain amount of dredging with the ‘‘ Agassiz’’ trawl
(see fig. 20) took place during the Scottish cruise. The
apparatus worked well, and some interesting hauls were
obtained. Figure 21 gives the appearance of a mixed

44 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

haul of Invertebrates when emptied on the deck.

Probably our most interesting capture with the dredge
was the Giant Sea-Pen (funiculina quadrangularis,
which was recorded* as follows in a letter which was sent
to “‘ Nature’’ from the “‘ Runa’’ on July 11th:—

‘‘Marine Biologists may be interested to hear that

Fie. 20. S. Y. ‘Runa’ with the ‘‘Agassiz”? dredge coming up at the stern.

the bed, near Oban, of the largest British Pennatulid
Funiculina quadrangularis, and the smaller Virgularia
mirabilis described by Mr. W. P. Marshall and the late
Professor Milnes Marshall, in 1881 or 1882 (I have no :
books of reference with me), is still apparently in very
flourishing condition. In a couple of hauls of the small
Agassiz trawl, from this yacht yesterday, between the
islands of Kerrera and Lismore, at depths of eighteen
to twenty fathoms, I got about a dozen fine specimens of

‘““NatuRe,” July 20th, 1911, p. 77.

MARINE BIOLOGICAL STATION AT PORT ERIN. 45

Funiculina, the largest of which measured nearly four
feet in length. The bed must be of considerable extent,
as the hauls were not on the same spot, and both
brought up equally good specimens of these magnificent
pennatulids. Most of the large specimens of Funiculina,

Fig. 21. Examining the Catch on deck.

by the way, were not caught in the trawl-net, but were
balanced across the front of the frame, at each end, in
such a precarious position as to make one wonder how
many others had been lost in hauling in. The bottom
deposit was evidently fine mud.”’

46 TRANSACTIONS LIVERFOOL BIOLOGICAL SOCIETY.

Tuer Microscoric LiFe oF THE BEACH.

An immense amount of work remains to be done in
examining with the microscope the various deposits, such
as sand and mud, found between tide-marks on our
shores—not once for all, but periodically; so as to
determine the nature of the minute animals and plants,
their relative abundance and their variations in quantity.
Some of these lowly organisms, although individually
insignificant, may exist in such quantities as to discolour
the sands or the sea-water, and even give rise to plagues
amongst shell-fish and other more valuable animals.
Invasions of this kind are known to have appeared in
America and in Australia, and a minute animal, hitherto
unnoticed in British seas, has been found repeatedly on
the tidal sands at Port Erin this year in considerable
quantity. I gave a preliminary account of this
occurrence to the Linnean Society of London on June Ist,
and described the later manifestations at the Portsmouth
meeting of the British Association in September. As
there have been some further changes since, I shall now
summarise the whole visitation, quoting some parts from
what was published by the Linnean Society in their
Journal,* and using, by kind permission of the Society,
the blocks which were prepared for that publication. The
matter began with the following observation :—

‘“In going to and fro between the village of Port
Erin and the Biological Station, during the recent Haster
vacation, those of us who were constantly at work had
occasion to take a short cut across the sandy beach at
least twice and sometimes six times in the day. One
gets into the habit, in these traverses, of looking closely
at the beach when the tide is out, on the chance of seeing

something of interest cast up. On April 7th, I noticed a
*Journ. Linn. Soc., Zool., XXXII, No. 212, p. 7].

MARINE BIOLOGICAL STATION AT PORT ERIN. AT

new and somewhat unusual appearance on the sand about
or a little above half-tide mark. The hollows of the
ripple-marks and other slight depressions formed by the
water draining off the beach were occupied or outlined by
a greenish-brown deposit which in places extended on to
the level parts so as to discolour patches of the sand (see

fig. 22).

rt ef hie

be iA he Pies : ii ci i

Fig. 22. The general appearance of the brown deposits in the ripple-marks
on the sand, reduced in size.

‘« Tn this position the deposit remained, more or less,
for a month—waxing and waning, sometimes increasing
in a tide, say, roughly tenfold, and at other times
apparently disappearing for a day or two and then
re-appearing either on the same part of the beach, or it
might be a few hundred yards away. At one time it
discoloured a continuous stretch of sand about fifty yards
long by five yards in breadth a little below high-water
mark, and was noticeable from some distance away.

“On first noticing it I supposed the appearance was
caused by a deposit of Diatoms, but on taking a sample
to the laboratory, microscopic examination showed
that although a few Diatoms (including Navicula
amphisbaena, or a closely allied form) were present, the

en
cr

48 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

deposit was formed almost wholly of enormous numbers
of a very active little Peridinian or Dinoflagellate of a
bright yellow colour (figs. 23 and 24). More careful
investigation, in which Mr. Riddell and Miss M.
Latarche helped me, enabled us to identify this form as
Amphidinum operculatum, described by Claparede and
Lachmann, in 1858, from specimens obtained at
Christiansand, Bergen, and a few other places in
Norway. |

es Ee: 3 (gt Sontee, Z

Fig. 23. Sand-grains and Amphi- Fig. 24. Part of Fig. 23 under
diniwm (photo-micrograph under high-power magnification.

low-power magnification).

‘The published records of Amphidiniwm, however,
do not give the impression that it 1s a common or
abundant organism. The latest comprehensive work on
such forms—the article on Peridiniales, by Paulsen, in
the ‘ Nordisches Plankton’ (Kiel, 1908)—recognises four
species of Amphidimum : A. crassum, A. rotundatum,
and A. longum, which as yet have been recorded from
Kiel only; and A. operculatum, which is stated to occur
in brackish water on the north coasts of Europe. In

MARINE BIOLOGICAL STATION AT PORT ERIN. 49

addition, Kofoid (‘ Dinoflagellata of the San Diego
Region,’ 1907) records A. lacustre from fresh water,
A. aculeatum, a pelagic form from Naples, and
A. suleatum, which he took in a vertical haul from ninety
fathoms in the Pacific. On hunting through the few
scattered references to A. operculatum which occur, one
finds, however, that R. S. Bergh, in the ‘ Zoologischer
Anzeiger’ for 1882, states (p. 693) that Spengel in
December and January found it in huge quantities on
the beach at Norderney. Although, therefore, Amphi-
dimum operculatum has been recorded once before as
occurring in quantity, the occurrence appears to be a
sufficiently rare event to be worthy of notice; and, so
far as I can ascertain, the species, although known from
several parts of North-west Hurope, has not been
previously found on the British coasts. I have written
to most of the marine laboratories (Plymouth,
Cullercoats, St. Andrews, and Millport) and to many
marine biologists, and have not been able to hear of any
British record.

“It is, however, not an unknown thing for rare
Dinoflagellates to appear suddenly in some locality on
an occasion in phenomenal quantities. Torrey, in the
“American Naturalist’ for 1902, describes the unusual
occurrence of a species of Gonyaulax on the coast of
California. Sherwood and Vinal Hdwards, in the
‘Bulletin of the United States Bureau of Fisheries’ for
1901, tell how for two weeks in September a Peridiniwm
infested Narragansett Bay in such numbers as to colour
the water blood-red and cause the death of many fishes.
Finally, Whitelegge, in the ‘ Records of the Australian
Museum’ for 1891, gives an interesting account of a new
species of Glenodinium (G. rubrum) which appeared in
such quantities in Port Jackson as to give the water ‘ the

50 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

appearance of blood’ and cause the death of great
numbers of oysters, mussels, and all forms of shore life.
Whitelegge supposed that the very heavy rainfall that
year, by affecting the salinity of the water, and then a
lengthened period of calm weather which followed, may
have provided favourable conditions for an unusual
development of the Dinoflagellata. The Glenodinium
appeared in vast numbers about the middle of March and
disappeared early in May. When it was at its climax,
the allied colourless species Gymnodinium spirale
appeared in the bay and soon increased greatly in
numbers and became finally even more abundant than the
red Glenodinium upon which it was evidently feeding.

‘* Returning now to our Amphidinium operculatum,
it is not easy to account for the sudden appearance of
this unusual Dinoflagellate (previously unrecorded in.
Britain) in such profusion on the beach at Port Erin last
April. Plankton hauls were being taken regularly across
the bay at the time, and they showed. no trace of the
organism. In fact, Amphidiniwm has not occurred in
any of the thousands of gatherings which we have taken
in the Irish Sea during the last five years, and which
have been examined in minute detail by Mr. Andrew
Scott.

‘Thinking it might be present in the shallow water
close to the edge of the beach, Mr. W. Riddell and I
took some hauls of the tow-net from a punt worked
backwards and forwards in a few inches of water as near
as we could get to the discoloured sand, but the
gathering, although it contained fine sand and mud,
showed no trace of our Dinoflagellate. It may be noted
here that although the size of the Amphidinium,
0°05 mm. in greatest diameter, is such that it can slip
through the mesh (averaging about 0°08 mm.) of the finest

MARINE BIOLOGICAL STATION AT PORT ERIN. 51

plankton silk (No. 20), still so much clogging of the
meshes always takes place in such hauls, and so many
other smaller organisms and particles of mud are
retained, that it is certain that had the Amphidinium
been present in any quantity in the water it would have
shown up in the gatherings.

‘““Careful scraping of the sand showed that the
Dinoflagellates were only in and on the surface-layer,
and therefore could not be regarded as coming up from
below. It occurred to us that possibly they might be
fresh-water forms derived from ‘the land; but we
ascertained that the little stream in the centre of the
bay, which in wet weather overflows on to the beach (at
other times it 1s conveyed into the town sewer), had not,
on account of the unusually dry season, sent any water
to the beach for some weeks. Moreover, on experimenting
with the hving Amphidiniwm in the laboratory, Miss
Latarche found that while it lived well in sea-water, or
when diluted with a little fresh, it died at once in fresh
and survived for a few days only in brackish water,
containing only a little sea-water. The exact salinities
of these mixtures were, unfortunately, not noted at the
time. Samples of the Amphidinium kept in shallow
dishes of wet sand at the Biological Station in a few
days showed such profuse growth that the sand was
covered by a dark-coloured layer, the water became
impure, and eventually all the Dinoflagellates died off.

“Observation under the microscope shows that
although this is a singularly active Dinoflagellate,
circling round and round with great vigour, so that a
drop of sand and water containing a number of the
organisms presents a most animated picture under a low
power magnification, still the Amphidinium seems to be
actually attracted to the sand-grains and associated with

52 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

them. The sand-grains in the field of view are always
peppered over with a number of specimens of the
Amphidimum (figs. 238, 24, 25), and if individuals be
watched they are seen after swimming round to come
back to rest on a sand-grain and remain there for a time
before starting off on another excursion. If they are
thus constantly associated with sand-grains or other solid
particles, and never swim more than a microscopic
distance from such a resting-place, that may account for
the fact that we have never found them in our plankton
gatherings.

Fig. 25. Sketch from living preparation, to show some Amphidinia resting
on the sand-grains and others swimming about (low power).

‘“ Amphidinium operculatum is also, however,
positively heliotropic, congregating in quantity on the
lighter side of the dish in the laboratory, and shifting in
bulk from the sand at the darker part of a tank to the
end nearer the window. This property accounts for the
invariable occurrence of the discoloured sand on the
surface only and never in the deeper layers.

‘“The published figures of this species are not very
good, so a view of both dorsal and ventral surfaces, as
seen under a high magnification, is given here (fig. 26).
There certainly seems to be a slight but definite cuticle
covering the greater part of the surface, although this

MARINE BIOLOGICAL STATION AT PORT ERIN. 53

has been denied by some previous writers. The two
characteristic Dinoflagellate grooves certainly join, as
is stated by Calkins, but not by other observers. The
_ posterior flagellum which projects freely from the body
: is not difficult to see, but the anterior one which lies
| along the transverse groove is not so easy to demonstrate,
and may differ a little in position and extent from what
is shown in the figure. Stages in longitudinal fission
were frequently seen, and that is probably the commonest
j method of reproduction. What appeared to be con-
jugation between two individuals was observed by Miss
Latarche in one instance.

—— — aeeeasrzremeasnyesnayitnstting -venctcmngttomnnt ranean ey
Se ha ea

Fig. 26. Dorsal and ventral views of Amphidiniwm operculatum—enlarged
from high-power magnification.

““TIt may be that this organism lives normally in
small quantities, so as not to be conspicuous, in some
region of the sandy beach, or possibly in some special

54 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

habitat beyond the beach, and that the present vast
increase in numbers has been due to some unusual
conjunction of circumstances; but what these were I am
not yet prepared to suggest. Several possible
explanations have occurred to us which we hope to test
by further observation. In the case of the Port Jackson
Glenodinium plague, Whitelegge thought the increase
may have been due to exceptional rainfall and calm
weather; but the occurrence this spring at Port Erin was
preceded by unusually dry and rather stormy weather.”

When giving this account of the matter to the
Linnean Society, on June Ist, I concluded by saying :—
‘“T am inclined to think that, although I can find no
previous record of such an occurrence, it is probable that
these swarms of Amphidinium have been seen before at
Port Erin, and possibly elsewhere. I fancy I have seen
the phenomenon myself in the past, and have supposed
it to be due to swarms of Diatoms, which certainly do
cause some of the yellowish-green and brownish-green
patches on the sand between tide-marks.’’ .

Two days after making this statement I was again
on the beach at Port Erin. I found in the same region
what was apparently the same patch of discoloured sand,
but on examining a scraping with the microscope found
that the deposit was now wholly composed of a golden-
yellow Naviculoid Diatom—one of the “ Amphisbaena
group’’. of Navicula (fig. 27), probably Navicula
(Calonets) amphisbaena, Bory. I searched the beach
carefully between tide-marks, and examined samples
from every suspected patch of sand, but could find no
trace of Amphidinium. The Navicula, which was present
in April in very small quantity (see above), seemed to
have completely replaced the Dinoflagellate. We have
probably still much to learn in regard to the comings and

MARINE BIOLOGICAL STATION AT PORT ERIN. 55

goings of such microscopic forms and their physiological
inter-relations in connection with what may be called
“the metabolism of the beach.”’

Fic. 27. Navicula amphisbaena. (?) Fia. 28. Navicula digito-radiata., (?)

For the photo-micrographs reproduced as Figures 13 to 18, 23, 24, 27 and 28,
we are indebted to our Hon. Treasurer, Mr. Edwin Thompson.

The Diatoms—both Navicula ‘‘ amphisbaena’’ and
another more slender form (fig. 28), very similar to
_Navicula digito-radiata (Greg.), along with a few
specimens of a larger form, apparently a species
of Pleurosigma—remained in possession of the
beach during most of June and July, and _ no
trace of the Dinoflagellates was seen for about
four months. But on returning to Port Erin, on
September 9th, after the meeting of the British
Association at Portsmouth, where I had made a state-
ment as to the April Amphidinium having been replaced

56 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

by the June Navicula, I found to my surprise. that the
brown patches on the sand were once more present, and
now showed dense swarms of Amphidinium, but no
Diatoms. This condition remained for a few days, and
then disappeared, and on the 16th the Naviculoid
Diatoms were back again in force and remained abundant
during the 17th and 18th of September. On the last two

dates some small and rather greener patches were found ~

which on examination proved to consist of a bright green
Euglenoid Infusorian. The Amphidinia made their
appearance next on October 2nd, and remained more or
less in evidence during the month. They were in great
abundance on the 12th and 13th, again on the 19th, and
finally, on the 25th and 26th. No Amphidima were
found between October 28th and November Ist,
but on November 2nd some small patches again made

their appearance in the usual positions on the beach and

then died away.

The following may be given as a brief tabular
statement of these remarkable alternations of the animal
(Amphidinium) and the plant (Diatoms) on the beach at
Port Erin during these seven months in 1911.

April 7to May1_ ss... Amphidinium, and a few Diatoms (Navicula)
June 3 to July 22... Diatoms (some Navicula, others Pleurosigma)
Sept. 9 and 10 ne Amphidinium in abundance, Diatoms absent
Sept. 16 to 18 “a Diatoms (naviculoid) a
Oct.2to 26... ae Amphidinium in abundance, Diatoms absent
Oct. 28 to Nov. 1... No Amphidinium present

November 2 ... ah Amphidinium (3 small patches)

Since this date neither organism has been found,
and no brown patches have been seen on the sand.

Presented in this form the evident alternation
between these two kinds of organism is very striking,
and, although it may be impossible as yet to give any
full explanation, still the facts seem to point to the
probability that the cause of the phenomenon is a
physiological one and that the explanation may consist in
showing that each organism in turn in its metabolism

MARINE BIOLOGICAL STATION AT PORT ERIN. 5)

exhausts or alters some essential constituent of the
environment so as to prevent its own continued existence,
in quantity, at that spot, but leaves the ground suitable,
or even favourable, to the physiological needs of the other
set of competing organisms.

L.M.B.C. Memorrs.

No further Memoirs since No. XIX, PotycH2#tT
Larvz (the young stages of the Higher Worms) at Port
Erin, by Mr. F. H. Gravely, M.Sc., have been published.
Buccrnum, the large whelk, by Dr. W. J. Dakin; and
Pacurus, the Hermit Crab, by Mr. H. G. Jackson, will
be ready for publication this winter; while Doris, the
Sea-lemon, by Sir Charles Eliot; Sacrrra, the Arrow-
worm, by Mr. Harvey; SapeLtarta, a_ tube-building
Annelid, by Mr. A. T. Watson, and other Memoirs are
also far advanced; and we hope to have a Memoir on our
Irish Sea Species of Ceratium and other Dinoflagellata
from Professor C. A. Kofoid, who did some work on the
local material during his visit to our laboratory in 1908.

The following shows a list of the Memoirs already
published or arranged for:

1 Ascrp1a, W. A. Herdman, 60 pp., 5 Pls.

II. Carpium, J. Johnstone, 92 pp., 7 Pls.

Ill. Ecuinus, H. C. Chadwick, 36 pp., 5 Pls.
TV. Copium, R. J. H. Gibson and H. Auld, 3 Pls.

VY. Atcyonium, S. J. Hickson, 30 pp., 3 Pls.

VI. LerrorutTueErrus anp Lernaa, A. Scott, 5 Pls.
VII. Linevs, R. C. Punnett, 40 pp., 4 Pls.
VIII. Puatce, F. J. Cole and J. Johnstone, 11 Pls.
IX. Cuonprvs, O. V. Darbishire, 50 pp., 7 Pls.
X. Paretua, J. R.A. Davis and H.J. Fleure, 4 Pls.
XI. Arenicoza, J. H. Ashworth, 126 pp., 8 Pls.
XII. Gammarus, M. Cussans, 55 pp., 4 Pls.
XIII. Anuripa, A. D. Imms, 107 pp., 8 Pls.
XIV. Liaia, C. G. Hewitt, 45 pp., 4 Pls.

58 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

XV. Antepon, H. C. Chadwick, 55 pp., 7 Pls.
XVI. Cancer, J. Pearson, 217 pp., 13 Pls.
XVII. Pectren, W. J. Dakin, 144 pp., 9 Pls.
XVIII. Exepone, A. Isgrove, 113 pp., 10 Pls.
XIX. Potycnartr Larva, F.H. Gravely, 79 pp., 4 Pls.
XX. Buccinum, W. J. Dakin. .
XXI. Pacurus, H. G. Jackson.
Doris, Sir Charles Eliot.
SABELLARIA, A. T. Watson.
SacitTa, E. J. W. Harvey. |
Cucumaria, KE. Hindle.
Oyster, W. A. Herdman and J. T. Jenkins.
Ostracop (CyTHERE), Andrew Scott.
Bueura, Laura R. Thornely.
Himantuatia, F'. J. Lewis.
Diatoms, F. E. Weiss.
Fucus, J. B. Farmer.
Botrytioiwes, W. A. Herdman.
Actinta, J. A. Clubb.
Hyproip, EK. T. Browne.
HALICHONDRIA AND Sycon, A. Dendy.
In addition to these, other Memoirs will be arranged
for, on suitable types, such as Pontobdella, a Cestode
and a Pycnogonid.

In addition to what can be recorded in this Annual
Report, there are some pieces of work which are
incomplete, or only begun, and there are many isolated
observations being. accumulated. Moreover, there are
Mr. Chadwick’s routine daily records of the physical
conditions of sea and air which may sometime prove of
interest in connection with plankton results and with
variations in the Fisheries. The diagram of Sea and —
Air Temperatures for 1911 is not yet completed, but
those for the two preceding years (which have not

ee
kr
s

MARINE BIOLOGICAL STATION AT PORT ERIN. 59

previously appeared in our Reports) are inserted here to
show the general similarity of the two curves, along with
a few points of divergence, and the manner in which
the temperature of the sea lags behind that of the air in
both winter and summer.

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PORT ERIN DURING THE YEARI9IO. 3

~ alee —_poypursyng 922469q
S gusuuapearghtneseseereenaseseesage

60 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

We append to this Report : —
(A) The usual Statement as to the constitution of the
L.M.B.C., and the Laboratory Regulations;

(B) The Hon. Treasurer’s Report, List of Subscribers, and
Balance Sheet.

APPENDIX A.

THE LIVERPOOL MARINE BIOLOGY ~
COMMITTEE (1911).

His ExceLttency THE Riagur Hon. Lorp Raauan, Lieut.- ~

Governor of the Isle of Man.

Rr. Hon. Sir Joun Brunner, Barr.

Pror. R. J. Harvey Grisson, M.A., F.L.8., Liverpool.

Mr. W. J. Hats, Liverpool.

Pror. W.A. Hervuan, D.Sc., F.R.S., F.L.S8., Liverpool.
Chairman of the L.M.B.C., and Hon. Director of the
Biological Station.

' Mr. P. M. C. Kermopz, Ramsey, Isle of Man.

Pror. BenzamMin Moore, Liverpool.

Siz CHartes Petrie, Liverpool.

- Mr. E. Tuomeson, Liverpool, Hon. Treasurer. |

Mr. A. O, WALKER, NALS 4) ietnce formerly of Chester.

Mr. Arnoutp T. Watson, F.L.S., Sheffield.

Curator of the Station—Mr. H. C. Cuapwicr, A.L.S.
Assistant—Mr. T. N. CREGEEN.

MARINE BIOLOGICAL STATION AT PORT ERIN. 61

CONSTITUTION OF THE L.M.B.C.
(Established March, 1885.)

I.—The Ossercr of the L.M.B.C. is to investigate the
Marine Fauna and Flora (and any related subjects such
as submarine geology and the physical condition of the
water) of Liverpool Bay and the neighbouring parts of
the Irish Sea and, if practicable, to establish and maintain
a Biological Station on some convenient part of the coast.

I1.—The Committse shall consist of not more than 12
and not less than 10 members, of whom 8 shall form a
quorum; and a meeting shall be called at least once a
year for the purpose of arranging the Annual Report,
passing the Treasurer’s accounts, and transacting any other
necessary business.

Iil—During the year the Arrairs of the Committee
shall be conducted by an How. Director, who shall be
Chairman of the Committee, and an Hon. TRHAsuRER,
both of whom shall be appointed at the Annual Meeting,
and shall be eligible for re-election.

TV.—Any Vacancres on the Committee, caused by
death or resignation, shall be filled by the election at
the Annual Meeting, of those who, by their work on the
Marine Biology of the district, or by their sympathy with
science, seem best fitted to help in advancing the work of
the Committee.

V.—The Exrensts of the investigations, of the publi-
cation of results, and of the maintenance of the Biological
Station shall be defrayed by the Committee, who, for this
purpose, shall ask for subscriptions or donations from the
public, and for grants from scientific funds.

VI.—The Brotoeican Srarion shall be used primarily
for the Exploring work of the Committee, and the
Specimens collected shall, so far as is necessary, be

62 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

placed in the first instance at the disposal of the members
of the Committee and other specialists who are reporting
upon groups of organisms; work places in the Biological
Station may, however, be rented by the week, month, or
year to students and others, and duplicate specimens
which, in the opinion of the Committee, can be spared
may be sold to museums and laboratories. :

LIVERPOOL MARINE BIOLOGICAL STATION
AT

PORT ERIN.

LABORATORY REGULATIONS.

I.—This Biological Station is under the control of the
Liverpool Marine Biological Committee, the executive of
which consists of the Hon. Director (Prof. Herdman,
F.R.S.) and the Hon. Treasurer (Mr. E. Thompson).

IT.—In the absence of the Director, and of all other
‘members of the Committee, the Station is under the
temporary control of the Resident Curator (Mr. H. C.
Chadwick), who will keep the keys, and will decide, in the
event of any difficulty, which places are to be occupied by
workers, and how the tanks, boats, collecting apparatus,
&c., are to be employed.
| Ili.—The Resident Curator will be ready at all
reasonabe hours and within reasonable limits to give
assistance to workers at the Station, and to do his best
to supply them with material for their investigations.

IV.—Visitors will be admitted, on payment of a small
specified charge, at fixed hours, to see the Aquarium and

A =
ee

MARINE BIOLOGICAL STATION AT PORT ERIN. 63

Museum adjoining the Station. Occasional public lectures
are given in the Institution by members of the Committee.

V.—tThose who are entitled to work in the Station,
when there is room, and after formal application to the
Director, are:—(1) Annual Subscribers of one guinea or
upwards to the funds (each guinea subscribed entitling to
the use of a work place for three weeks), and (2) others
who are not annual subscribers, but who pay the Treasurer
10s. per week for the accommodation and privileges.
Institutions, such as Universities and Museums, may
become subscribers in order that a work place may be at
the disposal of their students or staff for a certain period
annually ; a subscription of two guineas will secure a work
place for six weeks in the year, a subscription of five
guineas for four months, and a subscription of £10 for the
whole year.

VI.—Each worker is entitled to a work place opposite
a window in the Laboratory, and may make use of
the microscopes and other apparatus, and of the boats,
dredges, tow-nets, &c., so far as is compatible with
the claims of other workers, and with the routine work of
the Station.

VII.—Each worker will be allowed to use one pint of
methylated spirit per week free. Any further amount
required must be paid for. All dishes, jars, bottles, tubes,
and other glass may be used freely, but must not be
taken away from the Laboratory. Workers desirous of
making, preserving, or taking away collections of marine
animals and plants, can make special arrangements
with the Director or Treasurer in regard to bottles and
preservatives. Although workers in the Station are free
te make their own collections at Port Erin, it must be
clearly understood that (as in other Biological Stations)
no specimens must be taken for such purposes from the

64 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Laboratory stock, nor from the Aquarium tanks, nor from
the steam-boat dredging expeditions, as these specimens
are the property of the Committee. The specimens in
the Laboratory stock are preserved for sale, the animals
in the tanks are for the instruction of visitors to the
Aquarium, and as all the expenses of steam-boat dredging
expeditions are defrayed by the Committee, the specimens
obtained on these occasions must be retained by the
Committee (a) for the use of the specialists working at
the Fauna of Liverpool Bay, (6) to replenish the tanks,
and (c) to add to the stock of eae animals for sale
from the Laboratory.

VIII.—Each worker at the Station is expected to lay
a paper on some of his results—-or at least a short report
upon his work—before the Biological Society of Liverpool
during the current or the following session.

IX.—All subscriptions, payments, and other com-
munications relating to finance, should be sent to the
Hon. Treasurer. Applications for permission to work at
the Station, or for specimens, or any communications in
regard to the scientific work should be made to Professor
Herdman, F.RS., University, Liverpool.

MARINE BIOLOGICAL STATION AT PORT ERIN. 65

APPENDIX B.
mon. TREASURER’S STATEMENT.

In the following pages the Annual Subscription List
and Balance Sheet are shown.

Unfortunately the List of Subscribers is slightly less
than last year, which is much to be regretted, as the
expenses become greater year by year owing to the
increased work done at Port Erin.

~The Balance Sheet shows a small balance in hand,
but next year the expenses will be extremely heavy
owing to the publication of two new Memoirs, and more
funds for this purpose are badly needed.

Epwin THOMPSON,
Hon. Treasurer.

25, Sefton Drive.
Liverpool.

66 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

SUBSCRIBERS.

Beaumont, W. I., Citadel Hill, Plymouth
Briscoe, F. W., caine Isle of Man

Browne, Edward T., B.A. Saahiig: Berkhanite
Herts. :

Boyce, the late ‘Sir ater’ ERS, University
Liverpool

Brunner, Mond & Co., Rete icns.

Brunner, Rt. Hon, Sir John, Bate Silverlands,
Chertsey

Brunner, J. EF. L., MP, 28, ‘Weatherly Gardeag
London, S.W. sist

Brunner, Roscoe, Belmont Hall, cose
Bullen, Rev. R. Ashington, Heathside-road, Woking
Caton, Dr., 78, Rodney-street, Liverpool ...

Clubb, Dr. J. A., Public Museums, Liverpool
Cole, Prof., University College, Reading os
Crellin, John C., J.P., Andreas, I. of Man...
Crosfield, Harold G., Oxton, Birkenhead ...

Dale, Sir Alfred, University, Liverpool
Dixon-Nuttall, F. R., J.P., E.R.M.S., Prescot
Kliot, Sir Charles, University, Sheffield ..
Graveley, F. H., Indian Museum, Calcutta

balls NW sds 35, Lord-street, Liverpool ... a
Herdman, Prof., F.R.S., University, Liverpool ...
Hewitt, David B., J.P., Northwich |
iekson, Proi., i. R. S., University, Manchenem
Holland, Walter, Carnatic Hall, Mossley Hill
Holt, the late Alfred, Crofton, Aigburth, Liverpool
Holt, Dr. Alfred, Crofton, Aigburth, Liverpool
Holt, Mrs., Sudley, Mossley Hill, Liverpool

Holt, P. H., Croxteth-gate, Sefton-park, Liverpool
Isle of Man Natural History Society

Forward

DrENrPNFFRYF NF OFNPHF HOF OF HH EE 1p

£37

RSE f=) RS) ey RS Se Ss et

Sooo OS 2 SC ose Goo: oS 6 Gre esenue Sue

MARINE BIOLOGICAL STATION AT PORT ERIN.

Forward...
Jarmay, Gustav, Hartford, Cheshire se
Lever, Sir W. H., Thornton Hough, Cheshire ...
Lewis, Dr. W. B., W. Riding Asylum, Wakefield...
Livingston, Charles, 16, Brunswick-st., ries:
Manchester Microscopical Society...
Meade-King, R. R., Tower ea Pee
Mond, R., Sevenoaks, Keni..
Monks, F. W., Warrington..

Mosley, F. O., Woodside- fas eeeeeieal Park,
Huddersfield

Muspratt, Dr. E. K., Seaforth Hall, Bree ral
Narramore, W., peeehrides Avenue, Great Crosby

O’Connell, Dr. J. H., Dunloe, oe
Liverpool

Petrie, Sir Charles, Be onahines Chat ideo
Rae, Edward, Courthill, Birkenhead
Rathbone, Mrs. Theo., Backwood, Neston...

Rathbone, Miss eh Northumberland-street,
London

Rathbone, Mrs., ee a rice Tiveevael
Roberts, Mrs. Isaac, Thomery, 8. et M., France...
Robinson, Miss M. E., Holmfield, Aigburth, L'pool
Smith, A. T., 43, Castle-street, Liverpool...

Tate, Sir W. H., Woolton, Liverpool
Thompson & Capper, Manesty-buildings, Liv sthcel
Thomson, Dr. J. Stuart, University, Manchester
Thornely, Miss, Nunclose, Grassendale

Thornely, Miss L. R., Nunclose, Grassendale
Toll, J. M,, 49, Newsham-drive, Liverpool
Walker, Alfred O., Uleombe Place, Maidstone
Watson, A. T., Tapton-crescent Road, Sheffield ...
Whitley, Edward, Oxford ies

Forward

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68 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

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Weiss, Prof. F. EK., University, Manchester Le Wine
Wiglesworth, Dr., Rainhill.. 1 alae
Wragg, Sir W., D. ©. op Me Coyers St. Mary, Tele of Ae 1) gia
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SUBSCRIPTIONS FOR THE Hire oF ‘“‘ WorK-TABLBS.’’

Victoria University, Manchester ... ae | SOOM
University, Liverpool oe a ae A AO Tae
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REPORT on the Investigations carried on during 1911
im connection with the LancasHirE SEA-FISHERIES
Lasoratory at the University of Liverpool, and
the Sxa-Fish Hatcuery at Piel, near Barrow.

Drawn up by Professor W. A. Herpman, F.R.S., Honorary
Director of the Scientific Work; assisted by Mr.
ANDREW Scort, A.L.S., Resident Fisheries Assistant
at Piel; and Mr. James Jounstong, B.Sc., Fisheries
Assistant at the Liverpool Laboratory; and others.

(With plates and text figures.)

: CONTENTS. PAGE
1. Introduction (W. A. Herdman) - - - - = 71
2. Fish Hatching at Piel (A. Scott) - : = - = 78
3. Classes, Visitors, &c., at Piel (A. Scott) - - = = 81
4, Report on Plaice Measurements (J. Johnstone) - - 85
5. Internal Parasites and Diseased Conditions of Fishes,
with Plates I-V (J. Johnstone) - - 103
6. Report on Hydrographic Observations (Dr. H. Bassett) - 145
7. Onan Ulcerative Disease of the Plaice le Riddell and
D. M. Alexander, M.D.) - 155
8. On Public Health Bacteriology in the emcee Sea-
Fisheries District (W. A. *Herdman) - - 162
9. Bacteriology of Mussel Beds in Wyre (J. J ohnstone) - 187
10. Intensive Study of Plankton around South end of Isle
of Man—Part V (W. A. Herdman and A. Scott) - 197
11. Plankton of the West Coast of Scotland in relation to
the Irish Sea—Part II (W. A. Herdman and W. Riddell) 215
12. Note on the West Coast Lobster Fisheries Oe J. Travis
Jenkins) - 245
13. Appendix—Memoir on ie Hdible Whelk ‘(Buccinum
undatum) (Dr. W. J. Dakin) - - - - - 253
INTRODUCTION.

The account of the hatching of edible flat fish (plaice
and flounders) at our Piel establishment, the details of
which are given by Mr. Scott in the body of this Report,
shows that the work has proceeded on the normal lines
and has resulted, as usual, in about thirteen and a quarter
millions of young fish having been set free in the
Lancashire waters. To these have to be added the

i2 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

millions set free from the Port Erin Hatchery when
considering the Irish Sea as a whole. Care is taken in
the distribution from both hatcheries to return the fry
to the sea in suitable localities, such as are frequented
by the naturally produced fish of the same age. I desire
to point out, however, that these numbers must be
regarded as small, though it is difficult to see how, with
our present accommodation and organisation, they could
be substantially increased.

Mr. Scott also gives particulars of the practical
classes in Marine Biology and in Navigation for fisher-
men, and in Nature Study for school teachers, held in

the Piel Laboratory by Mr. Johnstone and himself ’

during Spring and early Summer.

Dr. Bassett, who for several years has kindly under-
taken the examination of the samples of sea-water
obtained in our hydrographic cruises, has now been
appointed Professor of Chemistry in University College,
Reading, but I am glad to say that he still continues to
carry on our work, and his report on the results obtained
during 1911 will be found printed below.

A great series of valuable statistics in regard to the
measurements of plaice, caught in the course of the
routine trawling experiments carried out by Captain
Wignall and the other Fishery Officers of the district,
is now being accumulated. An instalment of these,
bringing the matter up to date, is given by Mr.
Johnstone, but no general discussion of these data will
be attempted until a sufficient series has been accumu-
lated to render the conclusions independent of annual
variations. Mr. Johnstone, however, makes use of some
of these statistics as the basis for a short note on the
formula dealing with the relation of length to weight in
the plaice. This will be found at page 86.

OO OE

SEA-FISHERIES LABORATORY. 13

Fisu PARASITES AND FISH DISEASES.

Mr. Johnstone has an important article dealing with
certain new internal parasites of common fishes in the
district, and also with various diseased conditions in the
Ray, the Cod and the Flounder, which have been
investigated in the Laboratory during the year. All
these points are rather technical and cannot be briefly
summarised, but they all add important items to our
knowledge of the conditions under which our fish live.
A further paper on a similar subject, viz., a diseased
condition which affects the spawning Plaice in the
hatchery ponds at Port Erin, is given by Mr. W. Riddell
and Dr. Alexander, and the Bacteria found in the
diseased fish are discussed.

BACTERIOLOGY OF SHELL-IF'1suH BEDS.

The shell-fish question in its relation to Public
Health is becoming of increasing importance, almost year
by year. Mr. Johnstone, who carries out our bacterio-
logical investigations, has had further work in connection
with the Conway Mussel beds during the past year, and
has just made a report to the Fishery Board for
Scotland on the topographical and bacteriological con-
dition of the Oyster beds in the Firth of Forth, near
Edinburgh. On account of the growing importance of
this work, and the prospects of its increase in the near
future, I have thought it useful to give in the present
Report a summary and discussion of the bacteriological
investigations that have been undertaken in the past
under the direction of the Committee, and to point out
how necessary it is that the topographical relations of the
samples examined in the laboratory should be studied by
the naturalist in the field, Tides and other currents,

74 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

prevalent winds, shelter, depth of water, nature of
bottom and condition of the shell-fish bed, all affect the
distribution of the sewage and other organisms; and the
Bacteriologist who undertakes such work, unless he is a
Field-Naturalist and Fisheries expert and knows how to
allow for the various factors in the environment, runs
considerable risk of being deceived by the samples
examined and of arriving at erroneous conclusions as to
the condition of the shell-fish in relation to sewage
contamination.

PLANKTON INVESTIGATIONS.

Mr. Riddell and I have undertaken for a second year
a discussion of the plankton samples which I collected
from my yacht in the seas to the North and West of our
district. This year our cruise in July and August
extended from the Irish Sea to the Shetland Islands, so
we were able to sample undoubted Atlantic water of high
salinity bringing in Oceanic organisms. The relations
of the Oceanic water and its living contents to the periodic
changes and variations in the plankton of our coastal
waters are not easy to unravel, and our occasional
collecting expeditions to the Scottish waters north of this
district are undertaken in the hope of throwing lght on
the distribution of the plankton organisms and their
history throughout the year.

My investigations, with Mr. Andrew Scott, of the
plankton collected in the Irish Sea throughout the year
have followed the usual lines and are reported upon in
‘Intensive Study,’’ Part V, in the same manner as
before. The work is of such a detailed nature that no —
brief statement in regard to it can be usefully made.

SEA-FISHERIES LABORATORY.

~I
Or

FISHERIES EXHIBITION.

It was decided last Summer to arrange an Exhibition
illustrative of the fisheries of the district, and a grant of
money was made by the Committee in August for this
purpose. Most of the specimens required had already
been collected for the Fisheries Museum at Liverpool
University. Otherwise it would have been impossible to
have obtained them in the time at our disposal. Work
connected with this Fisheries Exhibit occupied a large
part of Mr. Johnstone’s and Mr. Scott’s time during the
Summer. Dr. Jenkins took pains to obtain for us some
fine specimens which were needed to complete the series.
My private Research-Assistant, Mr. Wm. Riddell, also
gave a good deal of time and valuable assistance in the
preparation of specimens, illustrations and labels for this
Exhibition.

I desire, however, to put on record for the informa-
tion of the Committee that, although a number of
different minds and hands have co-operated in the
preparations, this Fisheries Exhibition is mainly the
work of Mr. Johnstone, who has been indefatigable in
his efforts to obtain the best specimens and illustrations
and to arrange them in the most instructive manner.

In order to make the Exhibition more complete we
have had to include, on loan, some series (such as samples
of sea-bottoms, and of Plankton gatherings) that belong
to the Zoological Museum of the University, and some
results of work done at the Port Erin Biological Station
and elsewhere on the West Coast.

The collection was sent in the first instance, at the
request of the Fishmongers’ Company and the Natienal
Sea Fisheries Protection Association, to the Fisheries
Exhibition which was held in the Autumn at Rusholme,

76 TRANSACTIONS LIVERPOOL BZOLOGICAL SOCIETY.

Manchester. After that 1t was removed to the Public
Museum in Liverpool, where for some weeks it occupied
a very favourable position in the middle of one of the
large galleries, close to the Museum collection of British
Fishes; and there it was seen to great advantage and was
apparently much appreciated by the Liverpool public.
A special descriptive Guide with many illustrations was
prepared in order to put before the public the extent of

the Lancashire Sea Fisheries District, the Committee’s

administrative and scientific organisation, the nature of
their work, and some of their results. A number of
copies of this Guide were sold during the Liverpool visit.

In December the Exhibition was removed to the Art

Gallery at Oldham, and about the middle of January it
was taken to the Chadwick Museum at Bolton. Itis now
on the point of being moved to Preston, and arrange-
ments have been made for visits thereafter to several of
the other Lancashire boroughs. .

This summary-exposition, as our travelling Exhibit

may be called, of the application of science to sea-fisheries

investigation in our district ought to focus attention upon
what has been done during recent-years under the auspices
of the Lancashire and Western Sea-Fisheries Committee,
and should help in educating public opinion as to the
claims of such work for recognition and substantial
support.

The delay in receiving any response to the Com-
mittee’s application to the Treasury for a grant in aid
of such work, under the Development Act, has been most
disappointing and, from the point of view of the scientific
staff, almost disastrous. We are aware that the delay
has not been caused at the Treasury, nor by the Develop-
ment Commissioners. The pity is that during all this

time useful work remains undone for want of funds,-

—— ——

SEA-FISHERIES LABORATORY. 77

which we understand are waiting to be used; series of
observations at sea, which were stopped pending the
receipt of support from Government, cannot yet be
resumed, and the break in our records becomes monthly
more serious and may at some future time prove a fatal
obstacle to the completeness and validity of important
conclusions. In the interests of fishery research on the
West Coast, it is devoutly to be hoped that the subsidy
recommended by the Commissioners to the Committee’s
application may enable our full scheme of observations
and experiments to be resumed at an early date. It is
understood that the Commission has reported favourably
on our claims and that the answer from the Treasury may
now be received any day. We have suffered so much from
delay which we have been powerless to avert, that it may
be hoped that we shall now set an example to others by
promptly organising our scheme of work and expenditure,
in the event of an adequate grant being placed at our
disposal.*

I have placed as an Appendix, at the end of this
Report, a detailed memoir on the Edible Whelk
(Buccinum wndatum), prepared by Dr. W. J. Dakin, of
the Zoology department in the University of Liverpool.
The Whelk is of economic importance, both as a food-
matter and as a fisherman’s bait, and I am sure that
Fisheries Authorities and investigators will be glad to
have placed before them in this accessible form all the
information in regard to the animal’s structure, actions,
life-history and value that has been brought together by
Dr. Dakin. W. A. HerpMan,

Fisuerics LABORATORY,
UNIVERSITY OF LIVERPOOL:
March 25th, 1912.

* In Press, May Ist, 1912.—Since the above was printed notification
of a grant of £1,640 for the current year has been received, and the
new scheme of investigation under the Development Act has now
started. W.A.H.

718 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

FISH HATCHING AT PIEL.
By ANnpDREw Scott, A.L.S.

The results from the fish hatching carried on in the
spring of 1911 were almost similar to those obtained in
previous years. The adult plaice were trawled in the
autumn of 1910 in the closed area of Luce Bay by our
Fisheries steamer, and we have again to thank the
Fishery Board of Scotland for the necessary permission
to fish in this protected fishing ground. The flounders
were caught in the vicinity of Piel by the police cutter
stationed in the Northern section of the Lancashire
district. :

Unfertilised eggs from plaice and flounders were
noticed floating in the tanks for the first time on February
26th. Fertilised eggs from both species of fish were
collected and placed in the hatching boxes ten days later.
The spawning was earlier in 1911 than in 1910, and was
no doubt accelerated by the mildness of the winter.
Spawning at sea also appeared to be earlier in 1911 than
in the previous year, as many of the whiting caught by
the steamer on the off-shore fishing grounds near
Morecambe Bay light-vessel for dissection in the first
fishermen’s class were quite mature. When the fourth
class began on May 8th the fish in the tanks had practi-
cally finished spawning, and mature whiting were
unobtainable at sea. The spawning of the fish in the tanks
lasted two months. During that period fully one and a
quarter millions of plaice eggs were obtained, and
thirteen and a quarter millions of flounder eggs. The
incubation of the eggs was carried out in the Dannevig’
hatching apparatus, and the resulting fry afterwards
liberated in the sea.

The following tables give the number of eggs
collected and the fry set free on the dates specified : —

SEA-FISHERIES LABORATORY. 19

Puatce (Pleuronectes platessa, Linn.).

Eggs Collected. Fry Set Free.

March 8 ... 20,000 | L6,000= ... ~Apal 3
pee... 30,000 30,000... Bs 2
moe... 40,000 34,000... if 10
Seta 55,000 AOU | esas
mie... 65,000 DA OOS tack pct ae
eee... 710,000 65,500... ‘ ii
pee... 80,000 6F5000) |... 33 3
eee... 80,000 697500, =. 3 a
pees... 85,000 WOOSI era)
peo. 905000 Teg00, rp .

epee! ~... 95,000 BaroU0: 9 5, a
ap fee 30,000 (2p00— 2. May. — 1
a f= 80,000 GoFOM) re? 5 =
? a --.,' 79,000 69,500. ...: 7 se
i Pee. 15,000 6a,000 2: ‘ 8
wea ..- 65,000 D000)” :.0. ‘s F
foe... 00,000 AO OU tee: 6 ae
ig. 45,000 AO SU Does. me
~ ie... 30,000 24,000... " 18
» 25... 25,000 DO WOU rea. ee
feo... 10,000 S00. 5

May 2 ... 6,000 SOG: nu te
Total Eggs 1,271,000 1,100,000 Total Fry.

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

FiounpER (Pleuronectes flesus, Linn.).

March 8

Eggs Collected.

200,000
350,000
450,000
650,000
700,000
800,000
850,000
850,000
900,000
900,000
900,000
900,000
850,000
800,000
750,000
700,000
650,000
500,000
450,000
350,000
150,000

80,000

35,000

10,000

Total Eggs 13,775,000

Total Number of Eggs
Total Number of Fry

Fry Set Free.

176,000
300,000
388,000
579,500
600,000
710,000
707,000
757,000
800,000
800,000
800,000
800,000
757,000
710,000
662,500
600,000
579,000
442,500
388,000
300,000

~132,500

69,500
31,000
8,000

... March 27

' 12,148,000 Total Fry.

15,046,000
13,248,000

SEA-FISHERIES LABORATORY. 81

CLASSES, VISITORS, &c., AT PIEL.
By AnpReEw Scott, A.L.S.

Four classes for fishermen were held at Piel in the
spring of 1911. The Education Committee of the
Lancashire County Council voted the usual sum of money,
which enables forty-five fishermen residing in the
Administrative area to attend a course of instruction at
Piel. The Southport Education Committee sent four men
and the Blackpool Education Committee again sent three
men. The Liverpool Education Committee sent two
fishermen from the steam trawlers fishing out of Liverpool.
The Cumberland Education Committee, for the first time,
sent men belonging to that county. Altogether, fifty-
seven fishermen students attended the classes, and received
instruction in Elementary Marine Biology. ‘Twenty-
eight of them also received a course of instruction in
Navigation and Seamanship in addition to their Marine
Biology. The studentship holders were divided into four
classes—three of fourteen each, and one of fifteen men, as
shown by the following lists : —

Wiret Class, held March 13th to 24th—J. N.
Armstrong, Silloth; John Ferguson, Maryport; John
Butler, Flookburgh; James Hill, Flookburgh; Thomas
Cocking (Junr.), Morecambe; George Mount, Morecambe ;
James Cartmell, Blackpool; W. Gornall, Blackpool;
Jack Rimmer, Blackpool; Richard Howard, Southport ;
Benjamin Wright, Southport; John Wright, Southport ;
William Wright, Southport; Joseph Beck, Liverpool;
Frederick Houghton, Liverpool.

Second Class, held March 27th to April 7th.—Thomas
Pater, Whitehaven; William Wilson, Baicliffe; William
Hodgson, Flookburgh; Thomas Shaw, Flookburgh;

82 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Thomas Ghorst, Bolton-le-Sands; Arthur Townley,
Sunderland Point; Richard Bond, Morecambe; Thomas
Bond, Morecambe; Thomas Gerrard, Morecambe; Henry
Atkinson, Knott End, Fleetwood; John Abram, Banks;
Richard Abram, Banks; Richard Brookfield, Banks;
Thomas Abram, Banks.

Third Class, held April 24th to May 5th.—H.
Chapple, H. Crompton, R.J.Gornall, J. Harrison, F. Hill,
J. Johnson, A. Kissack, T. Nisbett, C. Price, J. Raweliff,
T. Singleton, S. Smith, (C. H.: Wilson) {Wo Vaden
Fleetwood.

_Fourth Class, held May 8th to 19th.—R. Cringle,
J. Cropper, R. Grundy, T. Hodgson, W. Holmes, J.
Huntington, R. Iddon, M. McMannus, T. Nisbett, W. P.
Sawyers, M. Sumner, W. A. Tennant, G. Wright,
J. Wright, Fleetwood.

In the first two classes the course of instruction
related to Marine Biology only, and was similar to what
has been given in former years. The third and fourth
classes were restricted to deep sea trawl fishermen, residing
in Fleetwood, who were preparing to sit for the Board of
Trade examinations for certificates. as second hand or
skipper of a fishing vessel. The morning lesson, lasting
two anda half hours, dealt with Marine Biology suitable
for deep sea fishermen. The afternoon lesson, lasting three
hours, was conducted by Captain E. Barker Thornber, the
County Navigation Instructor, who gave an efficient course
in Navigation and Seamanship. The continued success
of the Navigation courses has led to further development,
and it has been decided to told three classes for deep sea
trawlers in the spring of 1912. Only one class in Marine.
Biology will be held. |

Classes for first and second year courses in Nature
Study for school teachers were carried on betweer: April

SEA-FISHERIES LABORATORY. 83

26th and May 19th. These classes were again organised
by the Barrow Education Committee, with the permission
of the Chairman of the Sea-Fisheries Scientific Sub-
Committee.

The annual inspection of the classes by the Members
of the Sea-Fisheries Committee, and of the various Educa-
tion Committees of the County, was arranged to take place
on May 3rd. The day proved most unfavourable. The
party were unable to land from the steamer after she
arrived in the harbour, owing to a strong southerly
gale and heavy sea beating on the shore. A number of
representatives of the steam trawling industry at Fleet-
wood visited the class on May 11th, and were able to
see the men at work. Members of various rambling clubs
and a party of scholars from Barrow Secondary School
visited the establishment on the Saturday afternoons
during March and April. Mr. A. Harris, H.M. Inspector
of evening schools for the district paid an official visit,
and inspected the teaching work that was going on.
Mr. K. C. Dé, of the Indian Civil Service, came to see the
laboratory, along with Dr. Jenkins, in July. Dr. J. W.
Robertson, Chairman of the Royal Commission on Indus-
trial Training and Technical Education, Canada, paid a
visit to the establishment in September. He made
exhaustive enquiries regarding the methods employed at
Piel im carrying on the classes for the instruction of fisher-
men in Navigation and Elementary Marine Biology, and
the Nature Study (Marine Life) Classes for school teachers.
The equipment of the establishment was inspected, and
favourably commented upon. Mr. T. Kitahara, of the
Imperial Bureau of Fisheries, Japan, also called and
made inquiries about the work carried on. The following
letter has recently been received from the Director,
Department of Technical Education, Province of Nova

84 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Scotia, Canada. “‘ Will you kindly send me a full and
“detailed outline of the classes which you give in life
“history of fish, etc., for the special instruction of men
“engaged in the fishing industry. I note that you give
“the only class in England dealing with this very impor-
“tant question. It is of paramount importance in this
‘“ Province also, where the fishing industry is one of the
‘“three most important industries in the Province. We
“would lke to take some steps towards giving some
‘“adequate instruction to the men engaged in this
‘‘industry.—(Signed) F. H. Sexton, Director.’’

We have again to thank the United States Fisheries
Department; the Smithsonian Institution; Professor E.
Ehrenbaum, of the Biological Station at Heligoland ;
Dr. Annandale, Superintendent of the Indian Museum ;
Mr. E. W. L. Holt, the Scientific Adviser to the Irish
Fisheries Department; and others, for further additions

to our Library.

SEA-FISHERIES LABORATORY. 85

REPORT ON MEASUREMENTS OF PLAICE MADE
POEANG THE YEAR 1911.

By JAS. JOHNSTONE.

The measurements of plaice caught in the course of
the routine trawling experiments, carried out by
Captain Wignall and the [fishery Officers, have been
made in the usual manner. ‘They are recorded in the
following series of tables, and I do not propose to discuss
them here. For the most part the localities sampled are
the same as in previous years; and the statistics form a
-useful continuation of those already published. Captain
Wignall has, as in former years, devoted attention
chiefly to the summer plaice fishery near Nelson Buoy,
off the Estuary of the Ribble, and the winter fishery
near Great Orme’s Head, and in the adjacent bays, and
his series of figures for the last three years is sure to be
useful. Mr. G. Eccles gives a very complete series of
measurements of plaice caught near the Estuary of the
Mersey, particularly for a remarkable fishery which
occurred in 1911 near West Hoyle Bank, opposite the
Estuary of the Dee. This is just the sort of work that
is admirable: whenever such an exceptional abundance
of fish of any kind is observed it is most desirable that
the officers should make frequent hauls and obtain good
series of measurements.

Samples of plaice, sent by Captain Wignall and the
Fishery Officers, have also been examined as in former
years. Statistics of length, sex and age are given for
various fishing grounds. It is proposed to discuss these
when a sufficient mass of data has been obtained, and
it has become possible to deduce the general conditions
applicable to the principal fishing grounds, without
necessarily taking into account the variations from
year to year. Average weights for the various samples,

86 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and for each centimetre group have also been found, but
the tables are not published, since they can be
summarised in a future report. In this connection I add
a short note relative to the length-weight function in
the plaice.

The Length and Weight Relation in Plaice.

In 1903* Professor A. Meek showed that the weight in
ounces of a plaice is given by the relation w = al®,
1 being the length in inches and a a coefficient, which
was found in the series of figures considered to be about
00067. Subsequently Professor D’Arcy Thompson
suggested that the formula 3
(length in centimetres)*

100
should be used in fishery investigations. It was adopted
by Henking and Heincke in 1907,+ and has since been
generally employed.

weight in grams = k x

The coefficient £ ranges in value from about 077 to
about 12 according to the fishing ground, the season,
and the length of the fish. It is a reliable index of the
vague attribute known as the ‘‘ condition’’ of the fish.
When a plaice is plump and well nourished, and full of

roe, k is big; when it is thin, ‘‘ watery,”

or spent, k is

small. |
Generally speaking it is greatest in the early
summer months, and least in the late winter, since at the
latter season plaice usually cease to feed. In the case
of a summer fishery, such as that carried on during the
months, June to August, near Nelson Buoy and its
vicinity, this 1s the way in which / varies; but in the
case of a winter fishery, such as that carried on off the
coast of North Wales during the months of October to

**Northumberland Sea-Fish. Committee. Rept. on Sci. Inyests. for
1903. Newcastle, 1903, p. 40.

+ “Schollen u. Schollenfischerei,” Beteilig, Deutgchlands a.d. Int,
Meeresforsch., IV, V, Berlin 1907, ve |

SEA-FISHERIES LABORATORY. 87

January, é& attains its maximum value later in the year.
These remarks apply to immature fish and males. In
the case of mature females the development of the ovary
and the subsequent spawning produce variations which
have nothing to do with the question of the seasonal
changes of ‘‘ condition.’’

This coefficient & is the only convenient index of
condition. It is true that the average weight would be
just as useful, but we should then have to compare fish
of the same lengths. The coefficient can, however, be
found for an entire catch of fish, so that samples which
differ with respect to their canee of lengths can easily be
contrasted.

To find & we use the formula g = k—— i 100" putting it

in the form
100 g

> Ge)

g being the weight of all the fish in the catch, / the mean
length of each centimetre group, and f the frequency
at each mean length. The arithmetic involved is
laborious when, as is generally necessary, the fish are
arranged in centimetre groups; for the cube of each
mean length has to be found* and multiplied by the
number of specimens in the group, and the values found
have then to be summed.

But in the investigation of a catch of plaice, average
weights for each group are usually calculated in any
case. In the International Fishery Investigations the
lengths recorded are always means, thus all fish
between, say, n and m+ 1 cms. are recorded as
m5 cms. The graph of average weights is therefore a
a series of columns of base 1 cm. in

““histograph ”’
length, and the numbers are areas—the sum of the

88 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

average weights is the area of the figure between ia
graph and the z-axis.
If we call f(l) the length-weight function,

JOR
bh I f() dl is equal, therefore, to the sum of the average
=4il

weights. Now assume that f(l) is £l? and integrate this
expression for the range L, to L,. The coefficient x is

then easily calculated for
ieee 4 (sum of average weights)

(L.)*— (La)
Obviously it 1s necessary to add 0°5 to the highest

mean length to find Z,, and to subtract 0°5 from the
lowest mean length to find L,, the upper and lower limits
respectively.

Now, if such a series of average weights is found

and smoothed,’’ a curve can be drawn very

approximately through the points. If from the same

series the equation g = foe be calculated it will

generally be. found that its graph does not agree as
closely as it ought with the curve obtained by smoothing
the observed average weights. |
This suggests that the length-weight function
referred to above is not the best one. To find a better
one we employ the systematic ‘‘ method of moments ”’
used in biometric work, and assume that the series of
average weights is represented by the parabola
g=a+ bl 4 cl4+di2>+ ... Generally it is necessary
to find the constant a and the coefficients, b, c, d, and
to do this successive ““moments of inertia’’ must be
calculated from the rough statistics, and equated to
moments calculated from the theoretical equation. The
simultaneous equations so formed are solved to find the
constants. The method is clearly described, with
examples, in Palin Elderton’s ‘‘ Frequency Curves and

Correlation,’ and need not be further referred to here.

SEA-FISHERIES LABORATORY. 89

It is not laborious. It is true that the curve so
calculated may not differ greatly from that obtained by
the Meek-D’Arcy Thompson formula, but in some cases
it does differ sufficiently to render the latter formula
unsuitable for exact calculations. If, for instance, we
attempted to calculate the numbers of plaice above and
below a certain length (say the mean length at sexual
maturity) contained in a series of catches from a specified
fishing ground and season, from the commercial
statistics, we should have to find the length-frequency
equation, and the length-weight equation. I don’t think
there is any other way in which this could be done.
The ‘‘k formula’’ would in this case be unsatisfactory.

Generally speaking, I have found that a series of
average weights of plaice, from a definite ground and
season, can be represented by the equation

g=a+bl+ el?
if the series is a small one, i.e., the range of sizes
varying, for instance, from 14 cms. to 24 cms. With a
greater range another term may be necessary. But the
coefficiency of /° is always small and tends to vanish. It
may be negative, and in such a case extrapolation from
the curve is obviously unsafe.

But such an equation as is thus obtained would not
be nearly so useful as a means of comparison of the
condition of the fish, for all the coefficients would have
to be considered. Obviously the simpler formula is to
be preferred for such a purpose.

g = 83-21 + 12-207 + 0-636 12; or g = 0-97

100
g = 74:32 + 10-37 1 4- 0-109 12; or g = 0-96 ar
= 90-77 + 14-20 1 + 0-696 12; or g = 1-22 Se

94-33 + 13-81 1 + 0-697 12
139-15 + 17-70 1 + 0-649 1? — 0-0015 7s
= 147-13 + 19-20 1 + 0-934 1? + 0°0146 1s

The first term is, in all cases, the average weight
of the median group.

g
g
9
g

90 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

TABLES, I.—LENGTH FREQUENCIES.

Luce Bay, Off Shore from Morecambe Bay,
6 inch mesh, 1911. 6 inch mesh, 1911.
! September. March. April. July.
Mature Fish

16-5 57 = 1 = =
17:5 101 = 1 —_ 1
18-5 110 = 3 = 1
19-5 109 == 1 i 3
20-5 87 = 1 i 6
21-5 56 = 1 a 10
22-5 56 = 1 i 8
23-5 58 = 1 = 6
24:5 36 — 3 ] 6
25-5 50 a 2 1 4
26-5 45 = 1 2 1
27-5 44 = 1 = 5
28-5 39 = 1 = 3
29-5 67 as 1 — 2
30-5 58 1 1 2 1
31-5 59 3 1 as 1
32-5 80 4 1 2 =
33:5 62 13 — — ==
34:5 66 20 = a =
35-5 54 39 1 1 =
36-5 45 41 1 = =
37-5 29 29 = — —
38-5 25 25 —— = —
39-5 23 23 = — —
40-5 18 18 = = —
41-5 10 10 = = —
42-5 13 13 ~_ 1 =
43-5 8 8 = — -—
44:5 10 10 a“ 1 ua
45-5 6 6 =. i se
46-5 3 3 = — -—
47-5 4 4 = — a
48-5 1 1 2 1 =
49-5 1 i 1 = —
50-5 os = _- _ =
51-5 = oa — _ —
52-5 == = — -- =
53-5 1 1 os — —--

Totals 1491 Dal 3 25 15 58

SEA-FISHERIES LABORATORY.

Blackpool to Liverpool Bar, 6 inch trawl-mesh, 1911.

28-5
29-5
30-5
31-5
32-5
33-5
34-5
35°5
36-5
37-5
38-5

Totals 139

Nelson Buoy.

July.

w
co 2)
—

—
Sun! | |

—
CDW ODHE OP 10

—

99, TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Off Mersey Estuary, 6 inch mesh, 1941.

Horst CHANNEL, 6 inch mesh.

April. May. June. August. October.
10-5 — —— — — —
11-5 — 2 == = a
12-5 — a — — ==
13-5 1 9 7 eee =
14-5 -— 35 33 — ==
15-5 3 94 96 4 2
16-5 20 189 364 18 16
17-5 92 299 523 57 24
18-5 255 393 524 97 21
19-5 320 432 340 151 16
20-5 255 300 296 104 +
21:5 166 250 148 131 15
22-5 141 154 92 70 11
23-5 90 98 52 dt pal pe
24-5 61. 43 21 55 24
25:5 23 21 7 ratl 22
26-5 11 4 6 20 19
27-5 8 3 1 11 23
28-5 1 6 — 6 17
29-5 4 1 — 4 8
30-5 2 2 1 4 2
31-5 — 2 1 4 4
32:5 1 — — 2 3
33:5 1 — = — 2
34:5. — — = aes 1
35:5 — 1 = 2 1
36-5 1 — — 1 ae
37:5 1 — — — =
38-5 — — — — ==
39.5 — — = 1 1
40-5 _— — — = =
41-5 — — — = —z
42-5 — — — 3 =
43-5 — — — = eat
44-5 — —- — 1 =
45-5 — — — — =
46-5 — a = a ==

Totals 1457 2358 : 2512 818 257

SEA-FISHERIES LABORATORY. 93

Off Mersey Estuary, 6 inch mesh, 1911.

Orr West HoytEe Bank, 6 inch mesh.

Totals; 1470 5163 1041 | 398

June. July. August. |September.| October. | November.
10-5 — _- — — — —
11-5 = a= — — - —
12-5 _ — — — -— —
13-5 a= 3 -— — -= —
14-5 1 4 — ~ _ —
15-5 3 40 --- — — 2
16-5 19 129 zl 1 3 2
17-5 48 233 37 3 3 9
18-5 102 287 39 4 4 6
19-5 182 477 72 4 5 6
20-5 213 434 70 1 6 11
21-5 yh 521 88 3 5 7
22-5 165 612 134 14 6 18
23°5 139 601 102 21 3 31
24-5 143 529 119 28 5 40
25-5 93 506 135 42 10 69
26-5 78 295 100 54 12 77
27°5 60 209 57 67 8 91
28-5 25 129 36 60 19 96
29-5 21 74 18 50 24 89
30-5 6 39 13 24 10 88
31-5 _ 23 11 13 9 68
32-5 —_— 8 3 | 11 46
33-5 —- 9 -— 3 4 39
34:5 1 1 = 2 5 18
35-5 --- — — -- 3 7
36-5 — — | — —- 1 5
37-5 _- — — = 1 1
38°5 — — = pa = 1
39-5 — — _- — —— —
40-5 -— — — —- —, 1
41-5 — --- = — — —
42-5 —_— a —- — — =
43-5 —- — _ a om =
44-5 os — —- = — —
45-5 — —- — — — =
46-5 = — ~~ — — —_

157 | 828

94 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Near Mersey Estuary, Shrimp Trawl], 1911.

Jan.* | March.| April. | June. | July. Oct. Nov.
4-5 255 — 29 — 15 — 32
5:5 980 24 242 — 20 6 450
6-5 692 99 189 -— 1 39 1107
7-5 293 161 68 3 1 76 1475
8-5 161 322 40 5 15 5 725
9-5 lig 280 49 3 4] -- 217
0-5 46 176 61 2 41 1 1
1-5 34 2d 55 1 57 i 1
2°5 17 202 66 ] 32 — 3
3°5 13 188 45 2 10 — 1
4-5 8 149 81 — 9 1 Z
5-5 5 163 64 —. 6 1 =
6-5 3 106 74 i 11 — a
7:5 3 69 80 os 14 — =
8-5 2 77 49 —_ 18 — =
9-5 — 45 aii] — iy — i
0-5 — 31 28 — 20 — —
1-5 -- a 14 — 12 -— 1
2-5 _ 6 13 — 6 — —
3°5 == = + _— a — —
4-5 — 1 1 — — — —
5:5 a= 2 2 = a ua a
6-5 ae aie fs (ORES eae aus zt
7-5 —- — 1 — —— — —
urea aa Ee = 2s == ans as
9.5 imal eu com vere ook ee." ie
0-5 — — = — — — 1
31-5 — — — — — a iL
32:5 — _ — = — — —
33-5 — = — — — — —
Totals | 2683 2379 1292 18 | 360 | 130 4018

* Includes twe hauls, 28—30/12/1910.

SEA-FISHERIES LABORATORY. — 95

+ | Off Mersey Estuary, 7 inch trawl net, 1911.

January. March.

10-5 = —
11-5 ue eS,
12-5 — _
13-5 —- 1
14-5 -— 1
(15-5 mn Poa! 3
(16-5 3 10
17-5 13 14
18-5 23 24
19-5 26 41
20-5 26 25
(21-5 27 23
22-5 10 14
23-5 16 iby
«24-5 U7 14
25-5 13 1
26-5 9 —
- 27-5 8 —
28-5 3 —
29-5 1 —
30-5 -- —
31-5 1 —
32°5 — a
3-5 1 os
MOIS i. ... 198 * 188

96 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Conway, Beaumaris and Red Wharf Bays, 6 inch mesh, 1911.

)

10-5 — — — — — — — —
11-5 — — = == — — — —
12-5 — — — 2 — 1 — —
13-5 — ] —- 3 a 3 — —
14-5 — ae — 3 4 14 Lia —
15-5 6 ‘5 — 6 a 71 8 5
16-5 8 9 — 37 13 182 28 17
17-5 ey 19 — 24 34 280 61 21
18-5 10 10 2 28 31 253 37 18
19-5 21 6 — 29 18 - 206 46 20
20-5 13 5 i 19 25), | 129 18 23
21-5 19 4 18 13 17 101 25 19
22-5 14 6 22 8 10 96 19 16
23-5 14 6 lef 10 15 68 26 24
24-5 if oy 5) 16 9 8 47 ELS 15
25-5 i 6 13 9 11 44 19 32
26-5 6 10 6 it 39 17 29
27-5 3 4 8 5 13 42 12 18
28-5 5 3 3 L 5 46 a: 33
29:5 2 — 2 2 4 31 aL 16
30:5 -— — 1 4 4 18 12 26
31:5 — — — 1 1 24 12 18
32:5 —- 2 — 1 1 17 6 19
33:5 — — 1 — — 13 9 14
34:5 _- — — 1 — 11 6 13
35°5 — — — = = 3 4 8
36:5 treo — — — —_ a — 6
37-5 — nl — — — 2 3 1
38-5 1 1 1 — — — 1 2
39:5 — — — — — — 3 2
40-5 — = == = = — 2 1
41-5 — — a = — — 3
42-5 — — — = — — —_ 1
43-5 --- = = = == = — —
44-5 — = — = — : 1 —
45-5 1 — — — — sill —
46-5 -— —. -- -— — 1 — =
A7-5 —- — — — — — 1 —
48-5 — — = = = == = —
49-5 — — == = = == = —
50:5 — — = = = 1 = —
Tetals 147 99 121 220 230 1750 415 420

SEA-FISHERIES LABCRATORY.

Cardigan Bay, 6 inch mesh, 1911.

January. May. June.

= = 1

— 2 1

— 1 1

— 4 7

1 6 6

— 5 5

— 13 9

oe 14 11

2 13 6

3 19” 14

2 23 19

5 29 35

5 18 35

8 23 38

5 21 37

6 35 24

6 22 11

4 15 6

4 9 2

7 6 1

4 2 ——

1 2 _

3 2 ——

3 = fu

5 — =

3 3 =

1| — oe

2 ooo =

43-5 — _ —
44-5 1 — —
45-5 1 —- —
46-5 — _ —
47-5 2 -— —
48-5 — — —
49-5 — — —
50-5 — — —
51-5 — — —
52-5 — a —
53-5 2 -— ~
Totals 86 287 269

98 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Carnarvon Bay, 6 inch mesh, 1911.

| $s ———_—

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SEA-FISHERIES LABORATORY.

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Age Groups, off Mersey Estuary, 1911.

November, 1911.

SEA-FISHERIES LABORATORY. 101

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102 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

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SEA-FISHERIES LABORATORY. 103

INTERNAL PARASITES AND DISEASED
CONDITIONS OF FISHES.

By JAS. JOHNSTONE.

CONTENTS

COHNOMORPHUS LINGUATULA (VAN BENEDEN).
THTRARHYNCHUS BENEDENI (CreEty).
GYROCOTYLE URNA, GRUBE AND WAGENER.
GYRODACTYLUS ELEGANS, NoRDMANN.
MELANOTIC SARCOMA IN RAIA BATIS.
MELANOTIC SARCOMA IN RAIA CLAVATA.
FIBRO-SARCOMA FROM GADUS CALLARIAS.

LYMPHO-SARCOMA PRODUCING EXOPHTHALMOS IN
PLHURONECTES FLESUS.

9. ECTASIA OF THE SENSORY CANALS OF RAIA CLAVATA
WITH INTRA-CYSTIC MYXO-FIBROMATA.

10. CUTANEOUS PAPILLOMA FROM AHAIPPOGLOSSUS
VULGARIS.

PAPA OP

1. Coenomorphus linguatula (van Beneden).*

-I am indebted to a pupil of mine, Mr. Thomas
Newby, mate of the Fleetwood steam trawler ‘‘ Cygnet,”’
for four specimens of this most interesting Cestode.
They were taken from a Coalfish (Gadus virens) caught
to the north-west of St. Kilda, in 130 fathoms of water.
Mr. Newby had seen them frequently, but only in coal-
fish. He noticed that the worms were of a very unusual
appearance and preserved part of the liver of the host in
ice until the arrival of his vessel in port. Mr. T. R.
Bailey, Port Sanitary Inspector at Fleetwood, sent on
the specimen to me. All four worms were alive when

* The only references to the occurrence of this Cestode which I can
find are :—

1853. P. J. van Beneden, Pull. de l’Acad. Roy. Belgique, T. XX,
Partie II, p. 260, pl. I, Bruxelles.

1854. Diesing, Sitzwngsb. K. Akad. Wissensch. Wien, XIII, p. 591.

1889. Lonnberg, E. ‘ Ueber eine eigenthiimliehe Tetrarhynchiden -
larve. Bihang till K. Svenska Vet.-Akad. Handlingar. Bd. 15, afd. IV.
No. 7, pp. 1-48, pls. I-III. Stockholm.

104 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

received. They were adherent by their suckers and hooks
to the piece of liver, and two of them had excavated
cavities in the tissue. I tried to kill one in fresh water,
then in sea water contaming cocaine, but without success.
One specimen was preserved in Zenker’s fluid and serial
sections were made and stained with Mann’s methyl-
blue eosin. The rest were preserved in weak formalin.
Fixation and staining were quite satisfactory.

The appearance of the Cestode when alive is repre-
sented in fig. 2, Pl. I, about natural size. It varied
from 30 to 60 mm. in length, according to the degree
of contraction, for it was very mobile. When fully
extended it was quite smooth, without wrinkles or
furrows, a slight constriction marked off the anterior
‘“cephalic’’ part from the body, which tapered to a
blunt-pointed ‘‘ tail’’ extremity. When contracted after
killing, the whole body was irregularly segmented by
rather deep constrictions, and a short terminal portion—
the “appendix” of Lénnberg—was retracted into a socket
or sheath. In this condition the worm is figured by
the latter author, who also gives an account of its
morphology. Iam, however, able to add some details of
structure, and these may be worth recording since
Lonnberg’s paper is not easily accessible. At any rate,
the Cestode is so rare that confirmation of the already
published account may be desirable. |

The Scolex. In life the scolex is quite smooth, but
after fixation it is marked by a great number of
longitudinal, shallow furrows (Text-fig. 1). There are
two bothridia, ‘‘dorsal’’ and ‘‘ ventral’’ in position,
and each of these structures is an elongated suctorial
organ like the sucker of a Bothriocephalus. Its wall
consists of dense parenchymal tissue, with relatively few
muscle fibres. <A slight ridge runs longitudinally along

SEA-FISHERIES LABORATORY. 105

its floor, and the muscle fibres are much less prominent
along the median part of the basal wall than elsewhere.
These characters suggest that the bothridium really
consists of two structures, the adjacent walls of which
have fused together. It hes entirely below the general
surface of the scolex. Its posterior wall is entire, but
anteriorly the lateral walls thin out and disappear. The
proboscides, four in number, are situated at the anterior
extremity of the scolex. They are very short, almost
globular in form, and closely covered with short recurved
hooks. The dorsal and ventral pairs are in contact with
each other, but a little distance separates the two pairs.
Each proboscis is in relation to a sheath, into which it
may presumably be invaginated, though this did not
occur while I had the worms under observation. The
proboscis sheaths pass into muscular bulbs. All this
proboscidial apparatus resembles in every detail that of a
typical Tetrarhynchid, from which the scolex of
Coenomorphus differs only in the characters of the
bothridia.

Muscles of the Scolex. These differ in some respects
from those of the Tetrarhynchids. Lénnberg does not
give figures of their arrangement, so I have prepared
the diagram (Text-fig. 1). The muscle bundles originate
either in the proboscis sheaths, or in the walls of the
bothridia. The principal systems are:—(1) A very
compact bundle running ‘‘ dorso-ventrally ’’ between the
two bothridia, internal to the proboscis sheaths: this is
represented in fig. 3, Pl. I, in transverse section, and
diagrammatically in Text-fig. 1 by the darkly shaded
tract joiming the bothridial suckers. (2) Fibres origin-
ating in the axial parts of the proboscis sheaths. Other
fibres of this series, taking origin in the dorsal sheaths,
are inserted into both of the ventral sheaths, and vice

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

= eee

a Proboscidial nerve /
‘Lateral nerve cord Excretory canal

Fics. 1 and 2. Coenomorphus linguatula.

1. Diagrammatic transverse section through the Scolex showing the
arrangement of the muscles. The muscular parts of the bothridia and
the dorso-ventral muscle commissure connecting them are darkly
shaded. The muscle bundles radiating out towards the periphery are
represented as if they were projected in a single plane: in reality
they run obliquely. (See Pl. I, fig. 5, which is an actual representation
of the muscle fibres seen in one section some little distance behind the
transverse level taken as represented in the Text-figure). The numbers
of fibres drawn are in all cases less than those actually seen in the
sections.

2. The axial parts of the Scolex some distance behind the central
nervous system. The coarse connective tissue between the proboscis
sheaths; the proboscidial nerves, and the lateral nerve cords are
shown. The lateral nerve cord on the right side is seen giving off
branches to the bothridial suckers.

SEA-FISHERIES LABORATORY. 107

versa, an axial decussation being formed. The dorsal
sheaths themselves are not connected together by muscle
fibres, nor are the ventral ones. (3) Radial fibres inserted
into a peripheral fibrous zone beneath the integument.
Some of these originate on the outer parts of the proboscis
sheaths, while others seem to take origin in the dense
parenchymal tissue in the axial part of the scolex. These
bundles are represented as transverse in Text-fig. 1,
but they are really oblique (see fig. 5, Pl. I). They
are relatively strong and are very numerous. Not all the
fibres originating on the outer parts of the proboscis
sheaths belong to this series, for some of those starting
from the dorsal sheaths run towards the ventral parts of
the scolex, and vice versa, a decussation being formed on
either side. Others taking origin on the dorsal and
ventral parts of the sheaths run into the tissues of the
bothridial suckers. (4) Transverse fibres running across
from side to side of the scolex, and passing between the
proboscis sheaths. (5) Fibres passing outside the sheaths
in the lateral parts of the scolex; these run dorso-
ventrally. There are relatively few fibres im series (4)
and (5).

Series (1) and (2) are situated anteriorly to the
central nervous system. Behind the latter the only
muscle fibres are those connecting together the bothridia.
These (Text-fig. 2) run external to the proboscis sheaths.

Longitudinal muscles. These originate as two series
of fibres proceeding from the outer surfaces of the
proboscis bulbs. They become gathered up into two
sheets which (fig. 7, Pl. I) run backwards through the
appendix, dorsal and ventral to the main lateral excretory
eanals.. This arrangement is similar to that in other
Tetrarhynchids.

The Central Nervous System, This is represented in

108 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

fig. 3, Pl. I, which is based on a reconstruction from
serial sections. I am not at all sure that it is accurate
in all details, for the difficulty in the investigation of the
nervous system of Cestodes is that the fibrils stain only
with great difficulty, or not at all. What has stained in
the series of sections studied is the parenchyma, with the
ganglion cells, and it is these tissues that are represented
—of true nerve tracts there was no indubitable mdica-
tion in the preparations. But ganglion cells were
certainly present in the main cerebral mass, and some of
these bodies are represented in fig. 1, Pl. I. They
are typical bipolar or multipolar cells of variable size
with characteristic nuclei. They are usually situated in
spaces, the boundaries of which appear to be fine
reticula, with some nuclei. The processes of these
ganglion cells can be traced for a very short distance
only, and they appear to fray out into fine fibrils. The
tissue in which they are embedded is a modification of the
parenchyma, with a closer meshwork, and a rather more
intense staining reaction than elsewhere in the body.
Part of this ground tissue appears to consist of exceed-
ingly fine fibrils running in all directions, but whether
or not this 1s truly nervous and not neuroglial is difficult
to determine.

The central ganglionic mass hes immediately behind |

the strong dorso-ventral muscle bundle referred to above
as joining together the bothridial suckers. It is really a
commissural mass crossing the body from side to side, in
the middle line, and between the dorsal and ventral
proboscis sheaths (fig. 5, Pl. I). Four nerves take
origin, each by several “ roots,’ at the lateral anterior
margins of this ganglionic mass, and these—the both-

ridial nerves, in other Tetrarhynchids, run outwards and
forwards into the scolex.. Two large nervous strands take

SEA-FISHERIES LABORATORY. 109

origin from the lateral and posterior margins of the
ganglionic mass, and these run outwards and backwards
as the lateral nerve cords. At intervals branches proceed
from them into the tissues of the suckers. They are
elliptical in section and can be traced backwards as far as
the beginning of the appendix. Four other nerves take
origin from the anterior part of the central mass and these
(which are shewn in section, but not lettered, in fig. 5)
run backwards in pairs between the proboscis sheaths.
They become applied to the outer surfaces of the proboscis
bulbs and cannot be traced further. They are the
proboscidial nerves.

_ Distinct regions are indicated in the central nervous
mass, thus the anterior sections contain the ganghon
cells, while the posterior ones display only neurogiiai
tissues. The number of ganglion cells is quite small, all
could easily be reproduced in a plastic reconstruction of
the sections. The axial part of the mass appears to be
homogeneous, but the lateral parts appear to be differen-
tiated to form tracts running outwards towards the
various nerve roots. The proboscidial nerve roots can
easily be traced through the posterior sections of the mass
into the ganglionic regions.

The posterior regions of the mass consist mainly ofa
tissue which is certainly parenchymal in nature, and a
part of this is represented in fig. 4, Pl. I. I have
chosen a part which contains a ganglion cell. There is,
first of all, a framework of relatively coarse fibres or
trabeculae, bounding (in section) roughly polygonal
spaces. Filling this is the parenchyma, a tissue which
in sections appears to be a reticulum, but which, no
doubt, consists of homogeneous films meeting together so
as to include polyhedral spaces. In the preparations we
see, of course, mainly the sections of these films. Coarse

110 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

neuroglial fibres traverse this meshwork in various
directions, and here and there are nuclei, belonging to
the parenchymal tissue. Calcareous corpuscles, and
excretory capillaries are also present, but not so much in
the ganglionic region as in the more posterior parts.
This parenchymal tissue is, no doubt, traversed in all
directions by the hypothetical systems of nerve fibrillae
which pass out into the bothridial, proboscidial and
lateral nerve tracts. Of course, if it were not for the
presence of the highly characteristic ganglionic cells one
would have great hesitation in describing all this system
of parts as nervous. The form, it is true, 1s that of a
central nervous system and I have no doubt that such is
its nature. On the whole it appears to be simpler than
the corresponding series of parts in the Tetrarhynchids
described. |

The excretory canals. These conform in their
disposition to that of a true tetrarhynchid scolex. As a
rule, however, there are two main canals on each side of
the body; in the specimen described there is only one.
The branching and anastomoses of these canals is also
similar to the condition in allied forms; a peculiar
feature (noted-also by Lénnberg) is the presence of very
large sinuses in the posterior parts of the body: some of
these are represented in fig. 7, Pl. I. Fine excretory
capillaries are present everywhere in the scolex. Of
‘* flame-cells’’ I have seen no trace.

The Appendix. This is represented (in section) in
fig. 6, Pl. I. It is the terminal, conical extremity of
the body which is capable of retraction within a terminal
socket, or pouch, no doubt by the action of the longi-
tudinal muscles. It, and the adjacent part of the body
consists of a tissue rather different from that forming the

rest of the animal—not parenchymal in nature, but

—s

SEA-FISHERIES LABORATORY. ig lak

made up rather of very fine fibrillae running in various
directions. The lateral excretory canals approach each
other and finally fuse near the extremity of the appendix.
Loénnberg describes a terminal vesicle, but this is repre-
sented in my sections by the common part of the main
canal system and there is no real vesicle, only a cavity
having a stellate figure in sections. In close proximity
to the lateral canals, internal to them in the body but
external in the appendix, are two peculiar plexuses of
vessels. These are very narrow in calibre, have
relatively thick, homogeneous walls, and anastomose
with each other repeatedly. They appear to be excretory
capillaries. —

Nature of the Organism. Coenomorphus linguatula
is certainly a larval form, and it may be that it corre-
sponds to some adult Tetrarhynchid already described
from some large animal, such as a porpoise or shark
(since the host is itself a fairly large fish). But it differs
in several respects from the typical plerocercoid larva
of known species of Tetrarhynchus. The ‘cephalic
segment ’’ (bothridia and proboscidial parts) correspond,
but the ‘‘post-cephalic segment’’ differs greatly, thus
muscles and excretory canals are not present behind the
scolex, except in the walls of the receptaculum scolicis,
and then they form part only of the integumentary
system. The appendix, too, is something quite distinct.
What the organism suggests is the scolex and the
unsegmented “‘ neck’’ region of a Tetrarhynchid such as
T. erinaceus.

No trace of genital organs is present. Lonnberg, it is
true, describes the ‘‘ anlagen’’ of ovaries, testes and vasa
deferentia in the appendix of his specimen, but he gives
no figures, and I cannot help feeling that he has mistaken
the knots of excretory capillary vessels for these organs,

112 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Fixation plays strange tricks with the appearance of these
organs, and sometimes they are widely expanded, with
almost invisible walls, while at other times their lumina
are reduced to the merest chinks, and the walls may be
quite thick. :

The situation of the worm is also unusual. As a
rule a Tetrarhynchid larva inhabits the body cavity, but
it is enclosed in a cyst, derived partly from the larval,
partly from the host’s tissues. Coenomorphus, however,
lives freely in the peritoneal cavity attached by means of
its suckers and hooks in the manner of an ordinary
Cestode.

What we doubtless have here is a “‘ permanent ”’
larval stage. Gadus virens is, for Coenomorphus, a
collateral host, not a true intermediate host. I have argued
elsewhere* that this is the nature of the Teleostean
hosts of Tetrarhynchus erinaceus, which Cestode inhabits
only the Rays in its adult condition, but a number of
Teleosts in the plerocercoid stage. It is difficult to
believe that the Ray is infected by eating such fishes as
Gurnards and Whiting, in which fishes plerocercoid
larvae of 7. erinaceus are, in my experience, always
found. The true larval host is no doubt some small
invertebrate, a mollusc or crustacean, and both the
Teleosts and Elasmobranchs are infected by eating these
creatures. The plerocercoid and adult stages are, on this
view, collateral ones, as are the hosts. The same view is
also taken by Southwell with regard to the lfe history
of Tetrarhynchus unionifactor, which inhabits both
Teleosts and Elasmobranchs in Ceylon waters. But the
Teleost in this case is, according to Southwell, a cul-de-
sac in the life-history. |

Coenomorphus is therefore probably a Tetrarhy nehid

* Parasitology, Vol. IV, No. 4, January, 1912, p. 368,

SEA-FISHERIES LABORATORY. ce

larva which has failed to find its definitive adult host.
It is present in the (modified, no doubt) oncosphere
stage in some invertebrate which is eaten both by the
Coalfish and by the animal in which the Cestode 1s
sexually mature. Its situation is unusual, but so also is
that of 7. ervnaceus in the body muscles of the Megrim or
Halibut, and there, too, the larva is mobile, moving
about like a cheese maggot in its cavity; while it 1s
much larger than the larva of the same species which
inhabits the Whiting or Gurnard. The absence of the

larval cyst; the growth of a ‘‘neck’’ region; and the
direct attachment of the scolex to the tissues of the host,
are, however, features not presented by 7. er:naceus.
The reason that the sexual organs have not developed is
doubtless the absence of the specific stimulus to division
of the cell rudiments of these organs, afforded by the
fluids of the true adult. host.

‘Lénnberg suggests that his specimen might con-
ceivably have been a pathological form, but apparently
rejects this possibility. I have no doubt that it is not
pathological, and that the only departure from normality
is the capacity for an extended period of larval life, and
for greater growth than occurs when the regular life
history is experienced.

2. Tetrarhynchus benedeni (Crety).*

On July Ist, 1911, a local fisherman, working a
stake-net at Roosebeck, Morecambe Bay, caught 38
specimens of the Tope (Galeorhinus galeus), the fishes
varying in length from four feet six inches to five feet
six inches. All were females with well-developed
ovaries. One of these dogfishes was dissected by my

* Vaullegeard, A. ‘ Pecherches sur les Tétrarhynques.”’ Mem.
Soc. Linn, de Normandie, X1X* Vol. (ser. 2°, Vol. 3°) 3° fase. P. 265,
Pl. XIII. Caen, 1899,

114 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

colleague, Mr. A. Scott, at the Marine Laboratory, Piel,
and a single Cestode was found by him. I describe this
worm here. It is a Tetrarhynchus which I have not
previously seen; and its measurements are :—

Length of strobila: 85 mm.

Length of scolex: 1:92 mm.

Length of bothridia: 0°36 mm.

Breadth of bothridia: 0°42 mm.

Length of proboscis bulbs, 0°36 mm.

Length of terminal proglottis, 4:5 mm.

Breadth of terminal proglottis: 1 mm.

The worm is figured below. It is relatively long and
slender, more so than any other Tetrarhynchus which I
have seen. ‘

Scolex. 1, fig. 3. There are two bothridia, leaf-
shaped, more pointed anteriorly than at the posterior
end, where there is a shallow notch, as in the bothridium
of T. erinaceus. They project well out from the scolex,
and are relatively shallow. The proboscides, four in
number, arise from the antero-lateral margins of the
bothridia.

The remainder of the scolex—the ‘“ head-stalk’’—is
rather long, and decreases at first in diameter, swelling
out in the region of the proboscis bulbs, and thereafter
the diameter of the neck decreases greatly. The bulbs
themselves are relatively short.

These characters correspond fairly well with those of
Tetrarhynchus longicollis, and I thought at first that this
was the species represented. The armature of the
proboscides is, however, quite different, the hooks and
spines in each oblique row being different from each
other, as in the case of 7. ermnaceus. They are very
difficult to make out, but their general characters and
arrangement are represented in Text-fig. 3 (2 and 3).

SEA-FISHERIES LABORATORY. TS

The two views are those seen by focussing through the
same piece of a proboscis.

(a) There are two prominent and characteristic
spines in each oblique row, (2), which are long and
slender, and only slightly curved, and which generally
he across each other. At the base of each of these spines

q is a much smaller one with a wide slipper-shaped base,
and a sharply bent apical portion.

Fie. 3. Tetrarhynchus benedeni (Crety).

The Scolex and anterior part of ‘‘ neck”’ region,
Part of a proboscis.

The same part—the obverse view.

A bothridium seen en face.

The terminal proglottis.

(6) Opposite to these, on the other side of the
proboscis, is a group of three spines. One of these (.3) is
rather large, with expanded basal part, and sharply
curved apex. From underneath it there projects a

orm oo to

TENG TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

longer, slender, slightly curved spine; and close to it
there is another long spine, club-shaped at the base, and
sharply pointed at the apex.

(c) Between these two groups of spines are other two,
each group consisting of three. One of these is rather
long and nearly straight, except for its sharply bent tip.
The base is swollen. The other two are slender, gently
curved spines. }

All these groups of spines are arranged along the
length of the proboscis with great regularity ; and between
each two of the consecutive groups (c) there is a very
small, slightly curved spine which les nearly flat on the
surface of the proboscis.

These characters correspond fairly well to those of
T’. benedeni (Crety) as described by ae and I
identify the worm as this species.

The Proglottides.

Segmentation is very obscure for some considerable
distance behind the scolex. In the single specimen
obtained none of the proglottides is mature. The terminal
one is represented in Text-fig. 3 (5) and it will be seen
that it has the characteristic structure of a Tetrarhynchus
segment, resembling those of T. erinaceus very closely
except that it is relatively longer and narrower. It isin
the functional male phase, and the uterus is represented
only by a long narrow tube lying in the axial part of the
proglottis. The testicular follicles are very numerous,
and are rather small. The genital apertures are lateral
and may be situated on either side of the segment, there
being no definite order in their alternation.

3. Gyrocotyle urna, Grube and Wagener. |
Two specimens of this interesting Cestodarian were
found in the large intestine of a Chimera monstrosa,

SEA-FISHERIES LABORATORY. 117

caught off the south-west coast of Ireland and landed at
Fleetwood by a steam trawler. The worm has been
recently well figured by T. Scott,* and I have nothing
to add to his notes regarding its appearance and
occurrence.

4. Gyrodactylus elegans, Nordmann.

A number of specimens of a Trematode, certainly a
Gyrodactylus, and very probably the above species, were
observed on the fins of a small plaice, about 7 cms..
in length, ving in the aquarium tanks at the Marine
Laboratory at Piel. The fish was one of a lot that had
been trawled in Ulverston Channel, Morecambe Bay, but
though most of those retained in the tank were examined,
the Trematode was only seen on one specimen. Both
dorsal and ventral fins were infected, but not the tail fin,
nor the gills, and no parasites were observed on the
general surface of the body. ‘The fish could not be

Fic. 4. Gyrodactylus elegans. x 125 dia.

examined at once, and therefore the fins were cut off and
preserved in weak formalin. The state of the parasites
was far from satisfactory when they were afterwards
examined, and I have some doubt in identifying it as
G. elegans, though it seems to be very near this species.
The average length and breadth are about O'4 and
0°05 mm. respectively. The large sucker bears two long
recurved hooks, which are about 0°05 mm. in length; and
there are six or seven pairs of smaller hooks on the
margins, lateral and posterior. Most of the specimens
contained embryos.

*Twenty-eighth Annual Rept. Fishy. Bd. for Scotland, pt. IU,
pl. VIII, 1891.

118 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.
5. Melanotic Sarcomata in the Skate (Raza batis).

Two specimens of Skate containing such growths
were received during 1911. One was part of the “‘ wing,”
or pectoral fin, of a very large fish caught off the Blaskets
(County Kerry, Ireland) in 130 fathoms of depth. The
tumour was a very small one and was not sectioned.
The other specimen was also the wing of a Skate, caught
by a Fleetwood steam trawler when fishing off .Dubh
Artach Light (west coast of Scotland) in about 32 fathoms
depth. This specimen was received in a fresh condition,
and portions of the affected tissue were fixed in Zenker’s
fluid, in vom Rath’s fixative, and in Bouin’s fixative.
Sections were stained in iron haematoxylin, methyl-
blue-eosin, and by other methods. The fixation in
vom Rath’s fluid was the most satisfactory. In some
cases the tissues were decolorised by prolonged soaking
in hydrogen peroxide. The melanin is rendered colour-
less by this reagent, but the sections are very easily
detached from the slide during subsequent manipulation.

This latter tumour was a large, rather irregular
growth in the middle of the dorsal surface of the left
pectoral fin. It measured about 10 by 6 cms. and was
raised up above the general surface of the skin about
1 cm. Its surface was flat, but rather rubbed and
injured, and there was also some degree of general
softening, due to autolysis in the central parts of the
tumour. It was very soft everywhere, and was extremely
difficult to cut in the fresh condition. It was dense black
in colour, and the pigmentation extended for some
distance on to the adjacent parts of the integument.
There were several smaller growths on the rest of the
‘wing,’ and some of these were true metastases,
being distinctly raised up. Others were apparently only
pigment patches. One of the more obvious metastatic

SEA-FISHERIES LABORATORY. 119

srowths was cut out and sectioned. As a rule, these
secondary tumours and pigment spots measured about
lcm. in diameter.

There was no obvious indication of emaciation in the
parts of the fish seen by me or Mr. Bailey, who sent the
specimen.

One of the metastases sectioned, fig. 4, Pl. II,
shows clearly the locus of the growth. The epidermis
has gone from over the entire surface of the tumour and
is seen only at the edge, on the adjacent skin. Three or
four distinct layers of coarse connective tissue fibres are
seen in the section, and the cells of the neoplasm appear
to take origin in the finer areolar tissue between the more
superficial layer of these coarse fibres and the epidermis,
and in the deeper layers of loose connective tissue. The
darkly shaded region in the figure represents the
distribution of the sarcoma, and it is indicated by the
figure that it is initiated between the coarse connective
tissue bundles. These latter fibres themselves do not
grow or proliferate in any way, but they become broken
down by the growth between them of the cells of the
sarcoma, and probably suffer from lack of nutrition.
The lower layers of the dermis, the deeper areolar tissue,
and the structures included in it, nerves, blood vessels,
sensory canals and lymph vessels are not involved and
are normal in structure.

The same general distribution is seen in the sections
of the fully developed tumour, though the relations are
less easy to make out than in the metastasis. Furthest
away from the growing edge it is only the areolar tissue
directly underneath the epidermis that is involved: here
one sees the intrusive sarcomatous cells loaded with
melanin granules. Nearer to the centre of the tumour
all the fine tissue between the coarse fibres becomes

120 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

infiltrated, the fibres themselves being separated. Finally
the deeper dermal structures become involved, and in
the centre of the tumour the whole. integument has
broken down, but even there the underlying systemic
muscle bundles remain unaffected. |

Fig. 1, Pl. II, represents a typical part of the fully
developed tumour as seen under a high power lens. It
has been drawn from a section made from a part of the
tumour fixed in vom Rath’s fluid. The section was
treated with hydrogen peroxide, but the melanin was not
entirely removed. The stain was iron haematoxylin,
followed by eosin. ‘The sarcomatous cells are of various |
types. Many of them are spindle-shaped and two of
these are represented in fig. 2, Pl. II, isolated from
their surroundings. In some of these cells the nucleus
is situated at one end, giving the cell a club-like,
appearance, but usually it lies at about the middle of the
spindle. In these cells the nucleus is often quite normal
in appearance, though perhaps the chromatic skein, or
synapse, 1s more pronounced than in undifferentiated
cells, and the linin substance is colloid-like. As a rule,
however, the nucleus is not easily seen in these cells
and is, no doubt, the seat of melanin deposition and
degenerative changes.

Many other cells are short spindles, or are ‘“‘ oat-
shaped,’’ but the greater number are round, irregular,
and variable in size. Some of these are represented in
the figure; like the larger spindle cells, they possess
nuclei which vary very greatly in appearance, owing to
the amount of melanin deposited. No traces of nuclei
are to be seen in some of them, and the cell looks like a
locus only for the aggregation of melanin granules.

These cells, varying in shape and size from elongated
spindles about 0°08 mm. in diameter to round cells only

SEA-FISHERIES LABORATORY. 121

about one-tenth of this in length, with a very fine con-
nective tissue stroma, disintegrated cell fragments, and
apparently loose melanin granules make up the general
mass of the tumour.

In some parts of the tumour, and after successful
staining with methyl-blue-eosin, there are generally
distributed masses of ‘‘ eosinophilous’”’ cells, or numbers
of such cells lying singly among the surrounding
melanotic and connective tissue elements. Some of
these are represented in fig. 3, Pl. Il. They stain bright
red, and stand out clearly from the other cells containing
melanin. As a rule, their nuclei are not very easily
seen—the defect of the staining method, for treatment
with haematoxylin and eosin shows the nuclei clearly,
though the general distribution of the cells themselves
is not then so obvious. It is noticeable that many of
these brightly staining cells appear to be situated in
cavities, and to lie loosely with no obvious relation to the
stroma. For the most part they are either modified or
unmodified red blood corpuscles, and their general
distribution is due to the breakdown of the walls of the
capillaries and smaller arteries and veins. Fig. 3 thus
represents what is apparently a large capillary, or small
vessel, with an incomplete wall. Sarcomatous cells,
containing melanin, surround this wall, and it would
even appear that some of these cells adhere to it. Not
all the eosinophilous cells, however, are blood corpuscles,
some of them appear to be leucocytic in nature, and some
have deposits of melanin. This infiltration of the sarco-
matous tissue by migrant cells, or blood corpuscles, can
be traced in most parts of the tumour; in fig. 1, for
instance, a red blood corpuscle is shown.

The growth is, therefore, a melanotic, mixed-cell
sarcoma, showing a general tendency on the part of the

122 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

cells to assume a spindle shape. I have now seen four
examples of this condition, and Dr. G. H. Drew
records what is apparently the same thing. In all my
specimens the neoplasm is to all appearance what would
be described as a malignant sarcoma. In all it is chiefly
the upper layers of the dermis, more precisely the fine
areolar tissue, which is involved, and which has taken
on the characters of malignancy.

6. Melanotic Sarcoma in Ray (Raza clavata).

A large Ray, sent by Mr. T. R. Bailey, exhibited a
growth of this nature. The fish was about two feet in
breadth, and was landed at Fleetwood by a steam trawler.
It was in very poor condition. There was a large tumour
on the dorsal surface of the head at about the level of
the eye. The growth was nearly globular in shape,
measuring about 70 mm. in diameter. It was raised up
above the general level of the skin about 40 mm. At its
origin in the skin of the head its diameter was about
55mm. It was dense black in colour except in one small
area on the posterior surface. It was extremely soft,
damaged in one place, and almost semi-fluid below this
place of injury. Necrotic change had evidently taken
place. Where uninjured the tumour was covered by the
remains of the integument, and a few small spines were
present. There were a number of black patches on both
dorsal and ventral surfaces of the body. Some of these
were distinct metastases, presenting raised surfaces, but
most of them were pigment patches: they varied from
one to two cms. in diameter. The skin immediately
round the tumour was also pigmented. , nee.

A smaller tumour was attached to the posterior
border of the eye, projecting over the spiracle, the
opening of which was about half-covered by it. It was

SEA-FISHERIES LABORATORY. 123

attached by a narrow base to the integument covering
the eye, and no other part of the latter organ was
involved. It was yellow-white in colour, and very hard
and compact. |

The tumour on the head was very similar in appearance
to the melanotic spindle-celled sarcoma described by me
in last year’s Report*, but it was larger and softer, and
the metastases in the fish now described were not present
in the former specimen. In both cases there was
emaciation of the fish. In the tumour now being
described, diagnosis is more difficult than in the Port
Erin specimen, for necrosis, and secondary changes, have
made the tissue far less characteristic In appearance; and
owing to the overgrowth of the tumour itself on the
surrounding skin it is far less easy to trace the transitional
integumentary tissues. The neoplasm, however, involves
the connective tissues of the skin, mainly the layer
between the epidermis and the strong and coarse fibrous
bundles that lie underneath: this, and to a less extent,
the other connective tissue layers deeper down, are the
regions of active proliferation.

Fig. 5, Pl. II, represents a part of a section not far
from the growing region of the tumour. The fish had
necessarily been preserved in formalin so that the fixation
was not good. The sections were stained in various ways,
on the whole methyl-blue-eosin, or Ehrlich’s haema-
toxylin, followed by eosin, gave the best results. In the
older parts of the tumour the tissue elements are greatly
broken down, so that it is very difficult to identify them.
The whole tumour is deeply impregnated with melanin
granules, and these further obscure its structure, even
when much of the pigment has been removed by a

* Rept. Lancashire [Sea-Fish. Laby. Liverpool, for 1910; in Trans.
Lwerpool Biol, Soc., Vol.£25, 1911.

124 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

prolonged bath in hydrogen peroxide. Blood vessels are
fairly abundant, but their contents are generally difficult
to identify, and in many places the red corpuscles have —
broken down.

The basis of the neoplasm is a rather loose connective
tissue stroma, generally enclosing irregular, elongated
spaces. Apparently isolated nuclei lie among this
tissue: these evidently belong to the fibres. Within the
spaces of the stroma are cells which, on the whole, are
short spindles in shape. Many of them are loaded with
melanin granules, and in these cases the nuclear structure
is obscure: probably the chromatic skein has quite broken
down. In other cells, however, the nucleus has the usual
structure. Cell fragments, and even apparently loose
aggregations of melanin are also present, and the figure
shows a piece of one of the coarse connective tissue fibres
belonging to the layer which lies nearest to the epidermis.

The tumour on the eye is evidently a hard fibroma.
Part of it is represented in section in fig. 8, Pl. II, and
it will be seen that we have to deal with a tissue composed
almost entirely of relatively coarse fibrous bundles
running in all directions. The growth presents no very
remarkable characters.

7. Fibro-Sarcoma from Gadus callarias.

A growth cut from the snout of a cod, and sent to
me by Mr. Bailey, appears to be of the nature of a fibro-
sarcoma. It was about 75 cms. long, 4 cms. in width,
and was raised above the general surface of the head of
the fish about 3 cms. The outline figure below (Text-fig. 5)
represents its general appearance. It was greyish black
in colour; and it was not capsulated in any degree, its
tissue passing continuously into that of the head. It
showed two small areas of softening. It was received

SEA-FISHERIES LABORATORY. 125

preserved in formalin, and sections were made and
stained by Mallory’s connective tissue method, after
previous mordanting in Miiller’s fluid.

The tissue of the tumour was compact and hard, and
had all the appearance of a fibroma. But on examining
the sections it was seen that it was not a typical
fibromatous growth but possessed certain marks of
malignancy. Fig. 6, Pl. III, represents a part of the

LWWYd

Fig. 5. Cod with Fibro-sarcoma on snout.

tumour where the fibrous elements were mixed with
small round cells. Not all the tumour was so richly
cellular as this part, and in the denser parts relatively
few cells were present. In other places the cellular
elements were, however, much more abundant than in the
part represented by fig. 6. In fig. 7, for instance, part
of the tumour, consisting almost entirely of irregularly
shaped, somewhat stellate cells, is shown; and this may
pass into such a tissue as is represented in fig. 8, small
round cells, some showing a distinct tendency to become
spindle-shaped.

Such round-celled tissue, containing relatively little
fibrous elements, forms conspicuous nodules in the

126 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

tumour; and here and there are places where the fibres
themselves are seen to be undergoing a kind of colloidal
degeneration. 3
Over all the surface of the tumour the epidermis is
absent, and the superficial layers consist of coarse
connective tissue fibres.

8. Lympho-sarcoma producing Exophthalmos in a
Flounder (Pleuronectes flesus). |

The fish displaying this peculiar condition was sent
to me by Mr. T. R. Bailey. It had been caught in
Morecambe Bay, during August, 1911. It was a spent
female, 31 cms. in length, and in indifferent condition
with regard to nutrition. The pigmentation on the body
was normal, but the dorsal, ventral and tail fins were
almost white. The left eye was situated on the summit
of a protuberance about 21 mm. in height, and about
20 mm. in diameter at its base. Near the upper margin
this protuberance was very slightly constricted. It was
elliptical in section, measuring about 20 and 16 mm. in
its principal diameters. Seen from the ocular side of the
body, there was no trace of an eye, but on looking at the
protuberance from the blind side, the pupil could just be
distinguished. The cornea was nearly opaque, and.
through it the lens could just be seen. On cutting
through the cornea it was seen that anterior and posterior
chambers had fused: the iris was quite broken down;
and the lens was lying loose among a mass of leucocytes —
and blood corpuscles. The head of the fish is shewn,
two-thirds of its natural size, in fig. 6. The other
(right) eye was quite normal, and apart from the
exophthalmos, and the peculiar absence of colour on the
median fins, the fish was normal.

Text-fig. 7 ate ane a dissection of f the fish a

SEA-FISHERIES LABORATORY. 427

the blind side. The anterior part of the dorsal fin,
with its underlying body muscles and skeleton, has been
cut away, and the roof and left part of the cranium have
also been removed. The tumour is now seen to be more

Fic. 6. Flounder with Lympho-sarcoma. Nat. size.

extensive than its external appearance indicates. It fills
up the entire left orbit, forcing the eye out of its place.
All the eye muscles are present and are in nearly their
normal relations to the bulbus oculi. The inferior and
superior obliqui are not much different in position and
length from the same muscles in a normal fish: their

128 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

positions have, of course, led to the development of the
tumour in the part of the orbit behind them. The
internal and external recti are shown in the figure: they
are longer than in the normal condition, are attenuated,
and flattened out into sheet-like structures. The superior
and inferior recti are not represented in the figure, being

Rectus

jae
! ectus. |

crema!

Wi fi ; rior. WY _cChiasma \

=-=- i
Fe tery

: oe ae

hidden by the mass of the tumour. All the eye muscles
he on the surface of the tumour, and do not pass into
its substance at any place.

The optic chiasma is normal, and the right optic
nerve bends sharply upwards as it leaves the brain cavity,
and runs along the posterior surface of the tumour. It

really lies directly underneath the rectus externus, but

SEA-FISHERIES LABORATORY. 129

the latter has been pulled a little to one side in making
the dissection for the figure. The nerve itself is partly
atrophied. It is greatly flattened out, and near the
bulbus it consists of relatively few fibres. It cannot be
traced into the bulbus, and it is evidently undergoing
degeneration, although the imperfect fixation of the
specimen did not allow of this condition being minutely
studied.

Underneath the tumour, and between the latter and the
floor of the orbit, was a small flattened body, fatty and
semi-transparent in appearance, with a little black
pigment. This is indicated in fig. 7. It is apparently
the remains of the recessus orbitalis.

Nature of the Tumour.

The fish had been preserved in weak formalin before
being sent to me, and the fixation of the tumour was far
from being all that could be desired. Nevertheless,
almost all details of its minute structure could be deter-
mined, and I think there is little doubt as to its nature.
It was surrounded by a very loose investment of fibrous
tissue, and under a low power a complete transverse
section showed what appeared to be a number of bundles
of connective tissue radiating out from one main point
on the margin of the growth, with two other series of
bundles radiating out from adjacent parts of the margin.
Outside the tumour were the sections of the muscle
bundles, and that of the optic nerve.

The greater part of the substance of the tumour is
made up of loose, delicate, fibrous, connective or elastic
tissue bundles, running mainly parallel to each other, and
forming what might be called the trabeculae or frame-
work of a stroma, consisting of a very delicate reticulum.
But towards the external part of the tumour these fibres

130 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

are peculiarly modified: they become greatly enlarged,
or thickened, as if they were undergoing some kind of
colloidal degeneration. Fig. 6 of Plate II represents
such a thickening of the fibres, and their adhesion
together in bundles. These thickened fibres are at first
quite structureless, and they stain bright red with Mann’s
methyl-blue-eosin, and orange with Mallory’s connective
tissue stain. The large fibre in fig. 7, Pl. II, shows the
further progress of this degenerative change. The fibres
now display what appears to be a very delicate, faintly-
staining reticulum (blue with both of the stains men-
tioned), the interspaces of which contain a substance
which does not stain at all. Between these thickened
fibres is the general stroma of the tumour: this is
represented in fig. 7: it resembles nothing so much as
the reticulum of a lymph gland. This reticulum is
continuous with the enlarged fibres mentioned above.

All this reticulum contains great numbers of very
small cells occupying its interspaces, and two kinds of
cells are present: (1) small cells, the nuclei of which
measure about 0°002 uw in diameter. The cell bodies of
these hardly stain at all with either Mallory’s stain,
methyl-blue-eosin, haematoxylin and eosin, iron haema-
toxylin or Romanowsky’s stain. When the latter reagent
is apphed to a smear made from the tumour, the cell
bodies belonging to these nuclei can just be seen.
(2) Larger cells, staining red with Mallory’s combination,
and measuring about 0'005 mm. Some of these cells are
represented in fig. 1, Pl. HII. Im the parts of the
tumour where the connective tissue fibres are undergoing
the modification mentioned above, some of these cells can.
be seen (fig. 6, Pl. II) between the adhering bundles of
fibres, or even included in the structure formed by the
fusion of the latter.

SEA-FISHERIES LABORATORY. 131

Large spaces exist in all parts of the tumour, and
some of these are true lymphatic vessels. Usually they
contain the larger cells belonging to category (2), but
often they appear to be empty. Blood vessels are very
few and are difficult to identify. Some of the smaller
lymph vessels contain relatively large bodies, apparently
formed by the fusion of the cells mentioned, or by the
accumulation of some substance within them: the largest
of these bodies measures about 0°05 and 0°01 mm. along
its main diameters. ‘They are loaded with brown or black
pigment granules. Some are represented lying freely in
a lymph vessel in fig. 1, Pl. III.

Sometimes a small vessel, lymphatic or blood-
vascular, it is difficult to say which, contains numbers of
the cells (2) adhering to its walls. This suggests an
inflammatory process. |

In a section parallel to the main diameter of the
tumour, it is seen that the interior of the bulbus oculi
also contains the same kind of tissue that is found in the
tumour outside. The sclerotic is incomplete in the
section, so that a large cavity, much bigger than that
necessary for the passage of the optic nerve, must be
_present. There is no trace of the choroid layer, and only
the merest indication of the pigmented layer of the
retina. Through this cavity the foreign tissue is con-
tinuous. I think there is little doubt that the growth is
a lympho-sarcoma. It resembles strongly a small round-
celled sarcoma, but the connective tissue stroma is so
strongly developed, and the general suggestion of the
structure of a lymphatic gland is so striking that one
feels obliged to call it a lympho-sarcoma. The original
site of the growth was probably the choroid layer of the
retina. Identification of the growth as a glioma was
suggested by the destruction of the retina, but no traces

1382 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of glial elements could be made out in the tissues of the
tumour. There were no traces of similar growths in
other parts of the body, in the pseudobranch, for instance,
or in any part of the gill-system, so it does not appear
likely that the neoplastic cells were conveyed to the eye
in the blood stream from other parts of the body. Of
course, the peculiar vascular arrangements of the eye—
the ophthalmic artery originating in the pseudobranch,
and the choroid gland, suggested such an origin for the
tumour, but it is probable, I think, that the growth is a
primary one.

9. Ectasia of the Sensory Canals of Raia clavata with
intra-cystic Myxofibromata.

The head portion of a large Ray, sent to me by Mr.
Bailey, presented a most peculiar appearance on account
_ of the presence of large vesicles both on the dorsal and
ventral surfaces. On feeling these vesicles with the
finger 1t was apparent that they contained some solid
growths, but here and there they were soft, the walls were
thin and transparent and they contained only liquid.
Noticing the form of the vesicles more closely, it was seen
that they were symmetrically disposed on both sides of
the middle line of the head, and a curved line drawn
along their median parts corresponded very closely to the ©
direction of the sensory canals in a normal fish. The
vesicles are, in fact, the cephalic sensory canal system
dilated throughout its entire course, but with constrictions
here and there which confer on it the peculiar vesiculated
appearance.

Plate IV is the reproduction of a photograph of the
ventral surface of the head, and shows, on the right (of |
the photograph), the untouched condition of the canals, |
and on the left, the cavities cut open. If this is com-

SEA-FISHERIES LABORATORY. hoe

pared with a good figure of the sensory canals of the
Skate, Ewart and Mitchell’s for example*, it will be seen
that there is no doubt as to this interpretation. All the
canals figured in fig. 7, Pl. III. of the memoir cited are
present, though some are not clearly shown in the
photograph. That part of the infra-orbital canal lettered
1.0.4 to I.0.5 by Ewart and Mitchell; the part of the
hyomandibular lettered H.M.; the sub-orbital S.O.4 to
S.0.5; and the infra-orbital /.0.6 to 1.0.7 are greatly
dilated, and some of these dilatations present themselves
as vesicles about 35 cms. in diameter, and raised up above
the general level of the skin by as much as 1°5 cm.
Some of the canals are not much greater in calibre than
in the normal fish: these are the infra-orbital 1.0.7 to
I.0.8; the supra-orbital S.O.3 to S.O.4; the infra-orbital,
1.0.3 to I.0.4, and 1.0.6; and the hyomandibular
adjacent to its union with the infra-orbital. On a first
examination it appeared that some of the canals were
absent, but they were found beneath the floors of the
larger vesicles. This was the case with the hyomandi-
bular, and part of the supra-orbital. In these cases the
hidden canals were of the normal calibre.

There is not nearly the same amount of dilatation
of the canals on the dorsal surface of the head; but the
two supra-orbitals 7.0.7 to 7.0.8 are dilated, the greatest
diameter being about 15 cm. In front and external to
the left orbit there is also a large and complex cyst,
which is about 4 cms. in diameter, is raised up above
the general surface of the skin about 1 cm., and is
depressed below the surface about 1°55 cm. It occupies
the place of junction of the hyomandibular and infra-
orbital canals, and is almost certainly made up of

* Trans. Roy. Soc. Edinburgh, Vol. 37, 1891-2, pp. 87-105, pl. III,
fig. 7.

184 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

expanded portions of these tubes. From it runs forwards
a large canal, also dilated to about 0°5 cm. in diameter:
this is probably Ewart and Mitchell’s H.M.4 to H.M.8.
A smaller round cyst, about 1 cm. in diameter, projects
out from the antero-median surface of this large cyst,
and a large cyst, with semi-transparent walls, and about
25 cms. in diameter, lies on the floor. With these
exceptions, the dorsal surface of the head is normal.

The cavities contain a transparent, sticky fluid,
which is probably very similar to that secreted by the
sensory canal epithelia. Within them are the intra-
cystic growths, and these are very peculiar in nature.
All the dilated canals contain these growths, but in some
of them, the right supra-orbital on the dorsal surface,
for instance, the growths are small, round, worm-lke
bodies, slightly yellow in colour, and translucent. They
are smooth and are rarely branched or lobulated. Some
of these bodies are represented in fig. 2, Pl. IV.

The other growths are white in colour, fairly hard,
sometimes with a peculiar glistening appearance, and of
the most extraordinary shapes. Two of the larger ones
are represented in Pl. IV. The largest is about 8 cms.
in. length and about 1°5 cm. in diameter at its widest
part. The two growths represented were attached to the
internal wall of the cyst by very delicate pedicels, the
remains of which can be seen in the photograph, and
they were very easily detached. They were taken from
the expanded part of the infra-orbital canal, and one lay
in each of the large vesicles, but the larger of the two
projected into the hyomandibular canal. The other
dilatations on the ventral surface contained only the
smaller yellow bodies. It will be seen from the photo-
graph that the larger growths are produced into
appendages, lobes, and that they are racemose or

SEA-FISHERIES LABORATORY 185

botryoidal in form. It is difficult to describe them, but
the photographs convey a very good idea of their
appearance. The growths in the left supra-orbital canal
on the dorsal surface are quite similar, but part of the
floor of the cyst in this case is seen to be produced into
hard white ridges of varying form, and these can easily
be traced into the growth. Both in this canal, and in
the large cyst of the dorsal surface, the growths are very
firmly attached to the floor of the canal. In the large
cyst some of the growths have proliferated from the roof
of the cyst, and one of them has either broken through,
or has evaginated, so that it is visible without cutting
open the cyst. In the photograph of the ventral surface,
two of the growths are seen still attached to the floor of
the cyst.

Nature of the Tumours. Fig. 2, Pl. III, represents
part of a section of one of the smaller, white, irregular
growths, stained with Mallory’s combination. The
substance of the tumour is very uniform, consisting of
a fine fibrous tissue containing relatively few blood-
vessels. One of these blood-vessels is cut in the section :
it contains three red blood corpuscles and a leucocyte.
Outside the blood-vessel are a number of similar leuco-
cytes, and the aggregation of these cells round small
vessels is quite typical of the tissue. The remaining
elements of the tumour are very fine connective tissue
fibres, with very few nuclei. The fibres run in all
directions, except round the blood-vessels, where their
general course is concentric to the section of the vessel.
The tumours have a very distinct epithelium, continuous
with and similar to that lining the cysts. Its structure
(which is not easily made out on account of the formalin
fixation) is represented in fig. 3, Pl. III. The cells are
columnar, the free edges being usually confluent, or

136 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

apparently so; and the nuclei are situated usually about
their middles. Interpolated between them are elongated
granular bodies without nuclei, and coarsely granular,
and there are a few large ‘‘goblet’’ bodies. The
epithelium rests on a coarse reticulum, the exact
structure of which is difficult to make out.

Such a structure is quite consistent with the
interpretation of the tumours as intra-cystic myxo-
fibromata. They are apparently comparable in structure
with polypoid growths on the naso-pharyngeal epithe-
lium, and they are indeed present in a highly mucous
cavity. It is true that.the stellate nucleated cells with
long processes, described as present in typical growths
of this nature, could not be seen in the large white cysts,
but the latter were probably too highly developed, and
the typical structure need not, of course, be postulated
for identification.

The production of the cysts is, 1t seems probable,
to be accounted for by the occlusion, or the congenital
absence of the sensory canal pores. These are not very
numerous, and the absence of most of them would lead to
the production of mucus within the canals at a greater
_ rate than it could be removed. It was, of course,
dificult to be sure that the majority of these pores were
absent, but since the liquid in the large vesicles must
have been there under pressure, it is obvious that pores
could not have been present in normal number. There
may also have been occlusion of the canals at some places
by the intra-cystic growths, and it may well be the case
that the formation of these, together with the congenital
absence of pores, was the cause of the remarkable
dilatation of the canals.

SEA-FISHERIES LABORATORY. 137

10. Cutaneous Papilloma from a Halibut (Hippoglossus
vulgaris).

In November, 1911, Mr. F. Stokes, Port Sanitary
Inspector at Grimsby, sent me a piece of tissue weighing
about two pounds, cut from off the snout of a halibut
landed at Grimsby. The fish was about 120 lbs. in
weight, and the Inspector was of opinion that it was
‘“‘well-fed’’ and in good condition, apart, of course,
from the growth on the head. The latter, however, was
a very extraordinary one. It was very irregular in
shape, so that in the cut-out specimen it was almost
impossible to be sure of the relations of its parts to the
head of the fish. It was pigmented much in the same
manner as the skin of the upper surface of the fish, but
was perhaps darker in places. ‘The free surface was
everywhere thrown into fungoid, or ‘‘ cauliflower-
shaped ’’ excrescences. In some places these protuber-
ances were large, lobulated and botryoidal, presenting
in fact a great variety of appearances. In _ other
places the surface of the growth was very minutely
papillated, and dead grey white in appearance. It is
very difficult to describe the appearance of this growth.
It was very hard and dense, presenting in its in-
ternal parts all the appearance of a hard fibroid
tumour. In the deep the tissue was mainly aggregated
in nodular masses, presenting a dead white, sometimes
glistening, appearance. The tissue, both directly under-
neath the surface of the tumour and in the deeper parts,
was very difficult to manipulate when cutting sections.
It was extremely hard after embedding in paraffin, and
could only be cut with great difficulty. Staining, too,
was difficult on account of the formalin fixation, but
fairly good results were obtained with Hhrlich’s haema-

138 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

toxylin followed by eosin. I give, in the meantime, only
a provisional account of the structure of this growth.
Fig. 5, Pl. III, represents a vertical section through
the outer part of the finely papillated portion of the
tumour: the little protuberances standing out above the
surface are the sections of these smaller papillae. Below
them the tissue consists largely of bundles of coarse
connective fibres running in every direction. ‘Two series
of bundles do, however, assume prominence: (1) very
coarse fibres running nearly perpendicularly to the
surface of the tumour into the interior of the papillae,
and branching and apparently anastomosing freely;
(2) coarse fibres running approximately perpendicularly
to these in the deeper parts of the tumour. These also
branch, adhere together, and anastomose. In methyl-
blue-eosin all these coarse fibres stain at once a very
intense blue, and the same intense staining results from
treatment with Mallory’s combination. When treated
with carbol-gentian the lower parts of the tumour (at
this particular place) give a very decided mucin reaction.
Mingled with these coarse, intensely staining fibres
are finer ones, and these form a general stroma which

penetrates into the interiors of the papillae. Fig. 4,

Pl. III, represents part of the tissues within one of these
papillae, the external surface being to the left in the
figure. There is no evident epithelium on the surface of
the tumour, and no trace of epidermis, and one can only
see a kind of limiting membrane of very obscure
structure. The tissues within the papilla consist of a
fine areolar network with numerous nuclei, bundles of
coarser fibres, and some of the very coarse connective
tissue fibres from the lower parts—none of the latter is,
however, represented in the figure. The papilla is very
vascular, and sections of a capillary knot are shown:

SEA-FISHERIES LABORATORY. 139

these contain blood corpuscles, but the number of the
latter is relatively few. |

Everywhere in sections of the papillae, and in the
tissues directly underneath, are capillary tubes ramifying
to an extraordinary extent, branching and anastomosing
very freely. They are most numerous directly beneath
the surface, where they form an irregular layer, but they
are present also in the depths: they are represented in
fig. 5 by the lines of dots. Some of them are shown in
fig. 4, and are represented as filled with short fibrils. I
thought at first that the contents of these vessels were
granular, but examination under high power lenses shows
the fibrillar nature of this material. In unstained
sections this substance is coloured light brown in mass,
but where a few of the fibrils can be seen at the cut edges
of the preparation, they appear to be nearly colourless.
They present a glistening appearance when seen by
reflected light. They do not stain with methyl-blue-
eosin, Mallory’s combination, Ehrlich’s haematoxylin
and eosin stain, iron haematoxylin, or eosin alone. They
do not Gram-stain, nor do they take stain from carbol-
gentian. I thought at first that they might be bacterial
in nature, or perhaps the hyphae of a fungus, but the
negative staining reaction makes these interpretations
impossible. They do not react in any way to dilute
acetic acid, and cannot be crystals of lime. They are not
lipoid for they do not dissolve out under treatment with
xylol. They are some kind of inclusion in the tissues of
the growth, contained either in capillary blood-vessels or
in channels of their own.

The deeper parts of the growth vary in minute
structure. In many places what is seen is essentially
the condition already described in the case of the fibro-
sarcomatous tumour from the cod; that is, there is a

140 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

basis of either coarse or fine connective tissue fibres, and
included in the interstices of this are numerous small
round cells. Sometimes this tissue resembles that
shown in fig. 7, Pl. III, but it is always much denser,
and the cells are, relatively to the fibrous stroma, more
numerous. It is sometimes very vascular, but the
contents of the blood-vessels are peculiar. Sometimes a
smaller vein or artery contains practically unmodified
red blood corpuscles, but these are generally aggregated
together in the centre of the lumen of the vessel. In
other places the blood corpuscles appear as if they were
“clumped”? or agglutinated: the nuclei are few in
number, or entirely absent, and the cell margins are
indistinct as if the corpuscles had stuck together. Very
often the space between the axial mass of corpuscles and
the internal walls of the vessel is bridged by delicate
fibrils, radiating out in a stellate manner, and suggesting
the staining of fibrin filaments produced after intra-
vascular coagulation of the blood. I am uncertain
whether this coagulation has been produced in a natural
manner, or as the result of the fixation; practically
undiluted commercial formalin solution had _ been
employed for preservation. But I am inclined to think
that the intra-vascular coagulation is a natural reaction
produced in the development of the tumour. The walls
of the blood-vessels themselves are highly modified, and
sometimes cannot be distinguished from the surrounding
connective tissue stroma. If this alteration of the blood
has taken place as the result of some toxic substance
produced locally, it may be the case that the other
vessels, with their fibrillar inclusions, have also been
produced in this manner; that is, they may be capillary —
vessels containing crystalline products of the decomposi- ©
tion of the haemoglobin of the blood. I have already

SEA-FISHERIES LABORATORY. 141

seen extensive crystallisation of the haemoglobin in the
ease of a plaice which died in the tanks at Port Erin,
and which had ulcerated patches on its skin.

Elsewhere the growth has the characters already
described, with some modification. It is mostly fibrous,
sometimes finely fibrous, but at other times the structure
is very coarse and the fibres seem to be swollen and
degenerate, undergoing some kind of colloidal change.
Here and there are patches, or nodules of proliferation,
when there are very numerous small round cells. We
conclude that the growth is a cutaneous papilloma—a
kind of gigantic wart; with indications of a tendency
to the production of local malignancy.

EXPLANATION OF THE PLATES.
PxuateE I, figs. 1-7.
CoENOMORPHUS LINGUATULA (van Beneden). *

Fig. 1. Ganglionic part of the central nervous system.
Zeiss apochromatic 15 mm.

Fig. 2. The animal in the extended condition.
Shghtly reduced.

Fig. 3. The anterior part of the Scolex. Recon-
structed from serial sections, and seen from
the “dorsal” surface. Only two of the
proboscides, and two proboscidial bulbs and
sheaths are represented. Mag. about 10 dia.

Fig. 4. The central nervous system. The parenchy-
mal ground tissue is all that is represented
in addition to a ganglion cell and an excretory
canal. Zeiss apochromatic 15 mm.

* Tetrarhynchus megacephalus, Rud., may be the final form of
Coenomorphus.

142

Fig.

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Transverse section of the Scolex passing
through the central nervous system. Mag.
about 10 dia.

The figure should be compared with Text-
fig. 1, p. 106, which is based upon this and the
adjacent sections of the series.

Transverse section near the posterior extremity
of the animal, and passing through the
appendix. ‘The crescentic cavity is produced
by the involution of the integument con-
sequent on the retraction of the appendix.
Mag. 20 dia. ;

Transverse section passing through about the
middle of the animal. Mag. 14 dia.

Puate II, figs. 1-8.

Melanotic sarcoma from Raia batis. Round
and spindle cells containing melanin
granules; diffuse melanin granules; slight
connective tissue stroma; nuclei of cells either
modified or normal.

The same. Two typical spindle cells. Length
about 0°08 mm.

The same. Part of the tumour where a break-
down of the walls of the blood capillaries has
occurred. The darkly shaded cells belong to
the sarcoma; the lightly shaded ones are red
‘blood corpuscles.

The same. Section of a metastasis. The
darkly shaded part represents the sarcomatous
tissue; on the left, part of the epidermis. All
the structures shown belong to the integument.

Fig.

SEA-FISHERIES LABORATORY. 143

Melanotic sarcoma from Raza clavata. Cells
of rather irregular shape showing a tendency
to spindle formations. Marked connective
tissue stroma. The darkly shaded object is
part of a connective tissue fibre.
Lympho-sarcoma from the eye of Pleuronectes
flesus. Fibrous part of the growth, showing
colloidal swelling and fusions.

The same; an enlarged connective tissue fibre
showing a reticular structure. Fine meshed
reticulum enclosing leucocytes.

Fibroma growing on the integument of the eye
of Rava clavata. Coarse fibrous tissue with
few cells.

Puate III, figs. 1-8.
MoRBID HISTOLOGY OF FISHES.

Lympho-sarcoma from the eye of Pleuronectes
flesus. | Lymph space containing leucocytes,
some of which contain melanin. A smaller
lymph space on the right containing leucocytes
most of which are adherent to the walls.
Myxo-fibroma from Raza clavata. Fine fibrous
tissue running in all directions. A_blood-
vessel containing several blood corpuscles and
a leucocyte. Numerous leucocytes in the
tissue surrounding the vessel.

The epithelium covering the myxo-fibromata
sectioned in fig. 2. Columnar cells, with
interpolated mucus cells, resting on a coarse
connective tissue stroma, containing cells in
its interstices.

144 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Fig. 4. Cutaneous papilloma from the Halibut. Part
of one of the papillae represented in section
in fig. 5. Richly vascular connective tissue:
a capillary knot is cut through. Vessels
containing finely fibrillar inclusions.

Fig. 5. Cutaneous papilloma from the. Halibut.
Section of the external part of the growth.

Fig. 6. Fibro-sarcoma from the Cod. The more
fibrous part of the tumour, but containing
also numerous small round cells.

Fig. 7. The same. Irregular stellate cells.

Fig. 8. The same. Small round cells, some showing
spindle forming tendency. Relatively little
fibrous tissue.

Puate IV, figs. 1 and 2.

Fig. 1. Ray with ectasia of the sensory canals.
Ventral view of head. The canals are cut open
on the left, but are left untouched on the right
side. The intra-cystic growths have been
removed on the left side. |

Fig. 2. Dorsal view of the head. The canals are cut:
open, and the intra-cystic growths are present
am setu.

(Photos. by Mr. A. Scorr.)

Puce Vo feat l and ©

Fig. 1. Ray with ectasia of the sensory canals.
Photograph of two intra-cystic growths dis-
sected out from the canals cut open in fig. 1,
Pl. IV. Natural size.

Fig. 2. Papillomatous growths from the snout of a
| Halibut; about. one-half natural size.

(Photos. by Mr. A. Scort.)

Piate I.

Proboscis

Muscle
Lundle

LOscis:
Shaath

Ca/careous
corpuscle

71a

é Goer
Ves LF Central ganglionic
Ss

‘

lateral ‘Excréto

cana/

PHN peta, VND L
HARIRI ANH

AUNTY nvry f i, ry
est tg,
wr i vit wh ( "ei

- PRA! i

Longitudinal
-—~" muscles

ily ul
Wearle i

‘
at ig!
ta WK

Seger it [get eZ

F ey
5
4 ft

rake teb A ps

w. a “ —— TS aH roe |
ee LL ~ Ab ie ar fubips “a8, Live
teh? Ud i, rete ty
- pwn ad i Ptah teas tate i
Fig. ei 1 RTT PST WESTTLE TET ET POTS GST pe
AMULET ‘ hate i arg :

COENOMORPHUS LINGUATULA (van BeEneEveEN).

(J. J. del.)

MORBID HISTOLOGY OF FISHES.

see Se Saeed ae a ae ne Sy a ee ee eee

a eS a Se eS a nr

Prare BP

MORBID HISTOLOGY OF FISHES.

PEATE

IV.

(Photos. by A. Scott.) Fic. 1. Ventral aspect.

Fic. 2. Dorsal aspect,

RAY WITH ECTASIA OF THE SENSORY

CANALS.

PLaTeE V.

Fic. 1. Ray with Ectasia of the sensory canals. Two of the intra cystic growths.

Nearly natural size.

Pia. 2. Papillomatous growths from snout of halibut. Reduced about one-half.

DISEASED CONDITIONS OF FISHES.

(Photos, by A. Scott.)

“A
H

SEA-FISHERIES LABORATORY. 145

REPORT ON THE HYDROGRAPHIC WORK IN
THE IRISH SEA DURING 1911.

By Henry Bassett, Jun., D.Sc.,
Professor of Chemistry, Univ. College, Reading.

During 1911 considerable difficulty has _ been
experienced in carrying out the hydrographic work in
the Irish Sea. Samples were collected from the seven
stations on the lines, Piel Gas Buoy—Calf of Man, and
Calf of Man—Holyhead, on February Ist, June 12-13,
and October 24-25; and from Stations V, VI and VII
on December 10th. The June samples were collected as
a sort of compromise for those which should have been
collected in May and August, during which months
it was found impossible, for various reasons, to carry out
hydrographic cruises.

The observations made have, however, been
sufficient to show that the state of the water in our area
during 1911 was quite different from that found during
the two preceding years, 1909 and 1910, and more like that
found during 1907 and 1908 (with a probable difference
which will be referred to presently). Here, again, we
apparently have that intimate connection between the
salinities (that is to say, the state of the Gulf Stream
Drift) and meteorological conditions to which attention
has been drawn in the last two reports. It is hardly
necessary to point out that the brilliant dry summer of
1911 differed completely from the miserable wet ones of
1909 and 1910.

Table I, which summarises the salinities at the
Stations, V, VI and VII from the commencement of our
observations in July, 1906, is instructive. As has been
shown previously, these are the only ones of our Stations
which are affected by the Gulf Stream Drift—the eftect of

146 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the latter at the other Stations being entirely masked
by inflowing fresh water. Even Station VII, which is
nearest the Welsh coast, is liable to be affected by the
fresh coastal water, which probably accounts for the
somewhat irregular results at this Station.

On looking through Table I one is struck by the
remarkable similarity in the salinities at any particular
Station and at corresponding periods of the year during
the years 1907, 1908 and 1911. Stations V and VI are
particularly striking in this respect, while the results for
Station VII are somewhat less regular.

Similarly, the very close agreement of the salinities
throughout the two years, 1909 and 1910, is worthy of
particular note.

It certainly looks as though years when the salinities
are low during the winter months at these three hydro-
‘graphic stations (and probably at others as well), are
years when the following summer months are unusually
gloomy and wet. During such years it would appear
that the Gulf Stream Drift is so feeble that the maximum
salinity is not reached before May, and is then a good
deal lower than usual. In other years the maximum
salinity occurs several months earlier, and is a good deal
more pronounced.

In 1911, which was, of course, a quite abnormal year,
the maximum seems to have been reached at the very
beginning of the year or even at the end of December,
1910. I believe that this is quite unusual in our area,
and am inclined to associate it with the brilliant
character of the summer of 1911.

Unfortunately, we have no data for December 1906,
1907, 1908 and 1909, but from the general character of
the salinities during those years I believe that the
salinities during December were slightly lower than those

SEA-FISHERIES LABORATORY. 147

found during November. This is what we have found to
be the case in December, 1911, and it represents, I
believe, the more usual state of affairs. Since, moreover,
the high values of the salinities on February 14th, 1912,
at the three chief stations (Station V, 3465; Station VI,
34°47; Station VII, 34°38) indicate that they will almost
certainly prove to be the maximum values at these
stations for 1912, I have little hesitation in saying that
the summer of 1912 will probably be like neither the
brilliant dry one of 1911 nor the gloomy wet ones of 1909
and 1910, but just one of the somewhat variable and
uncertain summers which are usually experienced in
this country.

It is worthy of note that the salinities found during
February, 1912, are the highest we have observed since
commencing hydrographic observations in 1906. This
indicates the presence of an unusual amount of warm
water in the North Atlantic, and it is probable that the
wet and unsettled character of the winter and spring
months which have just passed is directly traceable to
this, for the presence of warm water is regarded by
hydrographers as favourable to the formation of cyclones.

Further work is still needed to show if the intimate
connection, which seems. to exist between the state of the
Gulf Stream Drift and the succeeding summer weather,
will hold over a long period. The question is so
important that it would be a great pity if anything
should prevent its accomplishment, and if it is not

possible for the Fisheries’ Steamer to collect the water
samples regularly, then some other arrangements ought
to be made.

Mr. J. Johnstone, B.Sc., collected the water
samples and made the temperature observations
as usual during 1911, while I have carried out the

148 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

salinity determinations as in previous years. The results
are collected in the tables, which follow, where T°, Cl°/,,,
S/o. and o, have the usual meanings. Monthly
means of daily sea temperatures at the surface, for the
years 1910 and 1911, are also given. These are calculated
from figures supplied by the Meteorological Office.

TaB.eE I.
Station. Year. Jan.- May. July- | Oct.- Dec.
Feb. Aug. Nov.
1906 — = 34-24 33°93 —
Vv 1907 34-40 34:33 34:13 34-00 —
53° 53° N. ;| 1908 34:27 34-25 34:05 34:05 —
4° 46’ W. a 1909 33°86 34-20 34-11 33°82 —
| 1910 33°86 34:07 34-00 33°84 34-43
1911 34:27 34-27 (Jun. 13) 34-09 34:00 |
1906 —- — 34-16 34:02 | —
VI 1907 34-33 34-29 34:09 33°95 —
53° 43’ N. 1908 34-40 34-14 34-14 34-18 —
4° 44’ W. 1909 33-93 34:22 34-11 33°95 —
1910 33°93 34-18 34-02 34:04 34-33
1911 34:36 34-23 (Jun. 13) | 34-14 34-05
1906 — — 34:13 34:04 —
VII 1907 33-75 33-96 33°98 33°73 —
53° 33’ N. 1908 34:38 34:05 34-14 34-02 —
4° 41’ W. 1909 33°98 34:14_| 34-00 33°84 — —
‘1910 33°73 34-13 33°89 34:04 34:16
1911 34-07 34:05 (Jun. 13) | 33-78 33-77
February 1, 1911.
Stations I. to IV. Surface observations only.
Station. Time. T°. 1} CPL cia yee ae

SE OO

T. 54°N.; 3°30’ W. | 10.45 a.m.| 5-0 | 17-67 | 31-92 | 25-26 —
Il. 54°N.; 3°47’ W. | 11.40 a.m.} 5-5 | 18-18 | 32-84 | 25-83
Tl. 54°N.; 4°4’ W. 12.30 p.m.| 6-6 | 18-66 | 33-71 | 26-48
IV. 54°N.; 4°20’ W. 1.30 p.m.| 7-0 | 18-85 | 34-05 | 26-69

SEA-FISHERIES LABORATORY. 149

Station V. (2.40 p.m.), 53° 53’ N.; 4° 46’ W. Depth of
Station, 43-9 metres.

Depth (metres)} T° Ol?/.,. aye ot
0 Cet 18-97 34-27 26-77
30 Heth 18-97 34-27 26-77
78 fers 18-97 34-27 26-77

Station VI. (3.55 p.m.), 53° 43’ N.; 4° 44’ W. Depth of
Station, 69-5 metres.

Depth (metres) Tr. tas cly

fefe) She Ot
0 7°75 19-02 34°36 26-83
30 T15 19-02 34°36 26-83
65 T-15 19-02 34°36 26-83

_ Station VII. (4.50 p.m.), 538° 33’ N.; 4°41’ W. Depth of
Station, 43-9 metres.

Depth (metres) Oley. wag Be ot
0 7-30 18-86 34:07 26-67
30 7:35 18-85 34-05 26-64
44 7:35 18-86 34-07 26-66

June 12 to 13, 1911.

Station I., 12/6/11 (4.5 p.m.), 54° N., 3° 30’ W. Depth of
Station, 23-8 metres.

Depth (metres) d ta Cl°/50 ae ot
sl, 15-1 18-03 32°57 24-10
22 11:3 18-28 33-03 25-21

150 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Station IT., 12/6/11 (6.40 p.m.), 54° N.; 3° AT’ W. Depth of
Station, 32 metres.

Depth (metres) sh C15. Dye ot
0 14-95 17-98 32-48 24-06

98 10-5 18-51 33-44. 25-67

Station TIT., 12/6/11 (7.30 p.m.), 54° N.; 4°4’W. Depth of
Station, 38-4 metres.

Depth (metres) re Cle/ 5. S'/ os Ot
0 14-2 18-34 33°13 24-72
35 10-95 18-66 33-71 25°73

Station IV., 12/6/11 (8.35 p.m.), 54°N.; 4° 20°W. Depth of
Station, 42:1 metres.

Depth (metres) ales Cee ee Ot
0 13-05 18-52 33-46 25-21
39 10-75 18-80 33:96 26-03

Station V., 13/6/11 (10.40 a.m.), 53°53’ N.; 4°46’ W. Depth
of Station, 82:4 metres.

Depth (metres) T° CV, So /er ot
0 10-2 18-97 34:27 26-37
30 9-7 18-96 34-25 26-44

70 9-7 18-96 34:25 26-44

SEA-FISHERIES LABORATORY.

Station VI., 13/6/11 (11.45 a.m.), 53° 43’ N.;
Depth of Station, 86 metres.

Depth (metres) al
0 10-8

30 10-35

70 10-35

C/o S"/o0
18-95 34-23
18-94 34-22
18-95 34-23

151

4° 44’ W.,

26-24
26-30
26-32

Station VII., 13/6/11 (12.35 p.m.), 53° 33’ N.; 4° 41’ W.
Depth of Station, 54-9 metres.

Depth (metres) sls
0 11-4

30 10-9

50 10-9

OV / 56 oy ae
18-85 34-05
18-84 34-04
18-85 34-05

October 24 to 25, 1911.

Stations I to IV., 24/10/11.

Station.

om. «DAN.
54° N.
54° N.
54° N.

we we we we

3° 30’ W.
3° 47’ W.
4°4’W.

4°20’ W.

of Station, 71 metres.

25-98
26-06
26-07

Surface observations only.

Time. A Laie Cs ek las i ea es
1.35 p.m. | 12-4 | 18-24 | 32-95 | 24-95
| 2.35 p.m. | 11-8 | 18-67 | 33-73 | 25-66
| 4.30 p.m. | 11-9 | 18-74 | 33-86 | 25-74
5.30 p.m. | 12-8 | 18-85 | 34-05 | 25-72
Station V., 25/10/11 (10 a.m.), 53°53’ N.; 4°46’ W. Depth
Depth (metres) jis Oe. 8/2 ot

0 12-85 18-87 34-09 25-74
30 12°75 _ — oe
66 12-75 18-87 34-09 25-76

152

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Station VI., 25/10/11 (11 a.m.), 53° 43’ N.; 4°44’ W. Depth

of Station, 73 metres.

Depth (metres) a
0 13:2

30 13-1

70 13-1

Cl’/.,.
18-90
18-90

S jee Of
34°14 25-53
34-14 25-73

Station VII., 25/10/11 (12 noon), 53° 33’ N.; 4° 41’ W.
Depth of Station, 62 metres.

Depth (metres) ur
0 13-3

30 13-2

58 13-2

18-70
18-70

See ope
33-78 25-41
33-78 25-43

December 10, 1911.

Stations V., VI., and VII. Surface observations only.

Station.

V: 53°53’N.; 4°46’W.
VI. 53°43’N.; 4°44’W.
VII. 53°33’/N.; 4°41’W.

Time.

1.4 p.m.
12.4 p.m.
11.4 a.m.

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154

SEA-FISHERIES LABORATORY. 155

NOTE ON AN ULCERATIVE DISEASE OF THE
PLAICE.

By W. Ripvpexz, M.A., Fisheries Assistant, Zoological
Laboratory, Liverpool ; |
AND
D. Moore ALtexanvER, M.D., School of Public Health,

Liverpool.

(With Two Plates.)

HISTORY AND CHARACTERISTICS,

For some years past the stock of spawning plaice at
the Port Erin Hatchery has been subject to a disease,
apparently infectious, which has done much damage.

The plaice are kept in two large open-air ponds,
which together occupy a space 90 feet long by 50 feet
wide, with a total capacity of about 130,000 gallons.
The number of fish in the ponds varies from about 300
to 400 or more; the average number is roughly 350.
Disease seems to have made its appearance first in 1905;
at any rate no diseased fish were observed before that
year. It has been more or less constantly present ever
since; in 1910 there was less disease than in any other
year since 1905, but in 1911 many of the fish in the pond
were affected.

The disease is characterised by superficial ulceration
(see Pl. I), which seems to have no very characteristic
site, though possibly ulcers are more common on the top
of the head and at the base of the tail. Still we have
seen ulcers on so many different parts of the surface that
we cannot regard any position as typical. The ulcers
are of a spreading and sloughing type, leading to con-
siderable destruction of tissue and often extending down
to the muscular layer. They vary in size up to about

156 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

3 cm. in diameter, with a well marked edge and a red
injected base. The lesion commences as a small inflamed
area of skin; this increases in size and ultimately the
skin breaks down to form an ulcer which gradually
enlarges. Asa rule more than one ulcer are present, but
they are never numerous. The fish dies eventually in an
emaciated condition, but infected fish have been observed
to feed freely, and even badly infected fish may retain
their muscular strength to a surprising extent.
Microscopically, a section through an ulcer reveals
great destruction of tissue, the base being covered with
débris. Beneath this are many swollen capillaries filled
with blood corpuscles, accounting for the injected
appearance which the base presents to the naked eye.
No bacteria were to be seen in the tissues. In the outer
layer of dead cells and débris were numerous slender
Gram-negative bacilli; these, however, we regard as
coming from outside, and probably not pathogenic.

BACTERIOLOGICAL INVESTIGATION.

The material for this investigation was derived
from : —

(1) A fish (I) found dead in the pond and sent to
Liverpool, examined when two days dead.
Cultures were made on to nutrient gelatine
from heart blood, liver, and from a large ulcer.

(2) Cultures were made by Dr. Dakin at Port Hrin
(A) from the ulcers of two living fish (II and

IIT) on gelatine, and (8) from the heart blood
and liver of the same two fish, also on gelatine.

All these cultures were at once despatched to —
Liverpool.
(3) A sample of water from the ponds.

SEA-FISHERIES LABORATORY. 157

Ulcers.
were examined. The gelatine cultures obtained were

Scrapings of four ulcers from three fish

diluted with normal saline and plated out upon gelatine
with a view to discovering whether the original cultures
were pure or a mixture of organisms.

From I a pure culture was obtained; the ulcers of
II and III produced a mixed culture of two organisms,
one of which was identical with that from I. All
liquefied gelatine within forty-eight hours.

Heart Blood.—F rom I a bacillus was obtained which
was identical with the bacillus of the ulcers. From II
was obtained a growth of a bacillus which was at once
differentiated from the others by its inability to liquefy
gelatine. No growth was obtained from III.

Liver.
from the ulcer and blood of the same fish. No growth
was obtained from II, and no culture was taken from the
liver of ITT.

Water.—Two distinct organisms were isolated. The
one closely resembled the organism obtained from the

I gave a bacillus identical with that derived

ulcer of I and one of the organisms from the mixed
cultures given by the ulcers of Il and III. The other
resembled the non-liquefying bacillus obtained from the
heart blood of II.

Three organisms therefore have to be described : —

Bacillus A: derived from the ulcers of all three fish,
and from the liver and heart blood of I. Upon agar this
gives a raised yellowish growth with clear transparent
edges. It grows well upon all ordinary solid media at
cold incubator temperature or at room temperature. At
37° C., in the warm incubator, the growth is scanty,
almost invisible on the few media upon which it exists,
and it loses its viability in two or three days at this tem-
perature. It has little or no action upon the common

158 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

sugars in 0'5 per cent. solution in peptone broth, and
milk is unchanged after seven days’ incubation.
Gelatine is liquefied in thirty-six hours. Morphologic-
ally it is a stout curved bacillus which shows longer
straight forms. It is Gram-negative.

- Bacillus B: derived from the ulcers of II and III,
and identical with the lquefying bacillus obtained from
the water sample.
This organism is closely allied to A in its character-
_ istics as regards growth at different temperatures. It
grows best at 20° C., less well at room temperature, and
apparently not at all at 837°C. Even at room tempera-
ture it loses its viability upon solid media in 5-6 days
and must be frequently sub-cultured. Its appearance on
solid media is very like that of B. colz, clear, transparent,
only becoming at all clouded at the bottom of the tube.
It liquefies gelatine within forty-eight hours. Morpho-
logically it is a slight slender bacillus, about the same
length as A, and also shows some tendency to curve. It
is Gram-negative.

Bacillus C. This differs from A and B in not
liquefying gelatine. It was derived from the heart
blood of II and from the sample of water. In shape and
size it resembles A, and is Gram-negative.

Smears were made from ulcer, liver, and blood of I,
and the accompanying micro-photographs (PI. II, figs.
1-3) show the distribution of the bacilli in these. Since
this fish had been dead at least two days when examined,
the tissue might have become invaded with intestinal
organisms and the bacilli seen may represent secondary
invaders. But it is significant that the cultures obtained
from this fish correspond with those from the others.
Further, no bacilli were observed in smears made from
liver and blood of two other fish, not ulcerated, which

SEA-FISHERIES LABORATORY. 159

were also sent to Liverpool after being found dead in the
pond. |

_ Bacillus A has some characters in common with
Bacillus Salmonis-pestis, which, however, is not viable
in sea-water. Whether any of these bacilli thus isolated
have an actual connection with the ulceration can only
be determined by experiments, such as those of Hume
Patterson on B. Salmonis-pestis, with fish ving in water
to which cultures of the organisms have been added. We
hope to carry out experiments of this nature with all three
bacilli and to incorporate the results in a future report.

CONCLUSIONS.

When the disease first appeared in 1905 Johnstone*
regarded it as due to an entomophthoran fungus, appar-
ently closely related to the genus Conidiobolus, which he
discovered in the viscera (liver, kidney, and mesenteries)
of some of the affected fish. He notes the characteristic
ulceration of the surface of the body. This fungus we
believe to have been a secondary condition. The fish
which died in 1905 were affected with the same super-
ficial ulceration as those which we have examined, and
this condition has been constantly present among the Port
Erin fish ever since. None of the fish which we have
seen have shown any signs of fungus in any of the viscera,
and it was by no means constantly present even in the fish
which Johnstone examined. He remarks that though
many of the dead fish showed no signs of fungus, the
surface lesions were of the same nature in all the fish.
He was unable to find any trace of fungus in the ulcers.
We believe, therefore, that this fungus was a secondary
condition, the case bemg comparable to that of salmon
disease. This fungus appears to have died out.

*Rep. Lanc. Sea-Fish. Lab. XIV, 1905 (1906), p. 179.

160 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Anderson* has recently described a very similar
disease among whiting and plaice from Bay of Nigg,
Aberdeen. He gives the following description of the ulcers
on the plaice:—‘‘ They commenced in paling and
desquamating spots, spreading rapidly to form large
ulcerated areas, often one to two inches in diameter.
The edges of the ulcers were deeply undermined, the base
often extending down to the muscular tissue. . . . The
base of the ulcers often presented a very injected appear-
ance.’’ This would apply equally well to the Port Erin
fish. :

The disease described by Anderson was apparently
more virulent than the present one, though its virulence
seems to have decreased towards the end of the period he
mentions; thus during September and October 187 dead
plaice were removed from the pond, while from that until
December 26th only 13 more were found.

Anderson regards the disease as some form of
septicaemic poisoning, possibly caused by sewage-borne
organisms. ‘The pathogenic organism seems to have been
the Staphylococcus pyogenes aureus, which was obtained
from all the superficial lesions-examined and from the
blood in most cases. In this respect, therefore, it differs
from the disease now described.

The Port Erin disease cannot be due to sewage

| pollution, as the water is remarkably pure, and the fish

are obtained from areas where there can be no question
of pollution. These septicaemic conditions may be due
to various organisms acting on fish which are under
abnormal conditions. As regards fresh-water fish,
Ceresole* has described a bacillus causing ulcerative
septicaemia in gold-fish (Carrassius auratus).

* 98th Ann. Rep. Fish. Bd. Scotland, Pt. IIT, 1911.
+ Zentr. fur Bakt. und Parasitenkunde, Bd. 28, 1900.

Prars. tl.

Ids 2.
DISEASED PLAICE.

BACILLI FROM DISEASED PLAICE,

SEA-FISHERIES LABORATORY. 161

It has been suggested that these lesions are due to
accidental injuries received either in the trawl or in the
storage pond. While it is not impossible that such
injuries may be a slight contributing factor, we cannot
regard them as the cause. If they were, it would be very
difficult to account for the absence of disease prior to
1905, though the conditions of capture and storage have
undergone no change since that year.

Taking all the evidence into consideration we
believe, though we do not consider it definitely proved as
yet, that this disease is bacterial and probably connected
with one of the three bacilli which we have described.
The condition is, then, a septicaemia strictly comparable
to those described by Anderson and Ceresole. All the
evidence appears to us to point to this conclusion.

EXPLANATION OF THE PLATES.

Puate I.

Fig. 1.—Head of diseased Plaice showing ulceration.
Fig. 2.—Another specimen showing ulceration at
base of dorsal fin.

Prats IT.
Fig. 1.—Smear from an ulcer of Fish I, showing the
Bacilli. From a micro-photograph.
Fig. 2.—Smear from liver of Fish I. From a
micro-photograph.
Fig. 3.—Blood-smear from Fish I. From a micro-
photograph.

162 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

PUBLIC HEALTH BACTERIOLOGY IN THE
LANCASHIRE SEA FISHERIES DISTRICT.

By Professor W. A. Herpman, D.Sc., F.R.S.

After twenty years of scientific work on shell-fish and
sewage in connection with the Lancashire Sea Fisheries
Committee, and just when the promised Government grant
will enable that work to be extended and put on a per-
manent basis for the future, it seems, now, the appropriate
time to summarise what has been done in the past and to
state the views and the actions which that work has led
up to. .

The Lancashire and Western Sea-Fisheries Committee
has jurisdiction over the largest shell-fish producing
areas in the British Isles; and the Scientific Staff of this
Fisheries District can claim to have been pioneers in the
application of scientific methods of research to these great
shell-fish beds, and especially in the investigation of
sewage pollution as a possible danger to the public health.
The connection between the consumption of polluted shell-
fish and epidemics of enteric disease is now too well
established to need further demonstration. It is admitted
in the Reports of the Local Government Board and of the
Royal Commission on Sewage Disposal and in many other
authoritative works. The chances of sewage pollution on
our populous shores are, and have been for the last few
decades, constantly increasing; and some sea-side
localities are certainly, in their present condition, quite
unfit for the cultivation or storage of shell-fish intended
for human food. On the other hand the magnitude of the
shell-fish industries around the British Islands, the
number of men and their families engaged directly or
indirectly, and the value of these food supplies to the

SEA-FISHERIES LABORATORY. 163

nation must not be forgotten. Here we have, on the one
hand a growing menace to the public health, and on the
other the threatened reduction, if not destruction, of a
great industry. The prospects of averting, or at any rate
of minimising, both these evils depend upon a more
intimate and accurate knowledge of the connection
between the shell-fish and the disease germs, and the
relation of both to their common environment; and the
opportunity has thus been given for scientific investiga-
tions on a large scale and leading to results of far-reaching
importance.
The local bacteriological work on shell-fish was begu

in the spring of 1895, when my late colleague Professor
(afterwards Sir Rubert) Boyce was visiting me at Port
Erin and we joined in work, both on the shore and in the
Biological Station, on the bacteriology of re-laid American
Oysters (obtained from Liverpool and Fleetwood) under
various conditions. These experiments showed that
oysters laid down only a short distance apart differed
enormously in their bacterial contents. Taking Bacillus
coli as an example, a certain standard culture made from
an oyster laid near the mouth of a small sewer gave
17,000 colonies, while a similar culture from those laid
a little distance off in purer water had only 10 colonies.
This work at Port Erin formed the subject of a paper read
by Professor Boyce and myself before Section D. (Zoology)
at the Ipswich meeting of the British Association in
September, 1895. Shortly after this, in October, 1895,
public attention was directed to the subject in a sensa-
tional manner by the serious outbreak of enteric fever
amongst those at the Stirling County ball, who had
unwisely supped on oysters which were afterwards proved
to be in-a very doubtful condition. This occurrence was
followed by a wide-spread ‘“‘oyster scare” which led to a

164 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

comprehensive investigation of shell-fish beds and layings
round the coasts of England and Wales, and the publica-
tion, in the following year (1896), of a Local Government
Board Report by the late Dr. H. Timbrell Bulstrode, with
an additional Report on the bacteriology by Dr. E. Klein.

In the meantime Professor Boyce and I continued
our investigations on the bacteriology of shell-fish under
various conditions, both at Port Erin and also in the
Zoological and the Pathological laboratories of the
University of Liverpool, with the help of several col-
leagues, and read successive reports at the Liverpool
(1896), the Toronto (1897) and the Bristol (1898) meetings
of the British Association. We also published a paper on
an unhealthy condition of American oysters associated
with the accumulation of copper in the leucocytes (Proc.
Royal Society, 1897); and finally incorporated all our
results in a larger work “Oysters and Disease,” issued as
a “‘ Lancashire Sea-Fisheries Memoir”? in 1899.

One of our first objects in all this work had been to
determine whether sewage bacteria, such as the Bacillus
coli communis and B. enteritidis sporogenes, occurred in
the alimentary canal of the living oyster taken fresh from
the beds—apart from what might be found in stored
oysters obtained in towns from markets and shops. We
also, in order to trace the history of the bacteria in the
shell-fish, infected oysters kept under experimental con-
ditions, and examined these after fixed intervals of time,
and so were able to show that the typhoid organism, for
example, could be recovered from our experimental
oysters up to 10 or 12 days after infection, and even under
some conditions up to three weeks from the sea-water
associated with the oysters.

In the 1896 report (British Association—Liverpool
meeting) we dealt mainly with the bacteriology of the

SEA-FISHERIES LABORATORY. 165

oyster and the behaviour of Bacillus typhosus in sea-water
and in the body of the shell-fish, and although we found
that we were, on occasions, able to recover our
experimental bacilli up to the twenty-first day from
infected sea-water kept in the cold, still in most cases they
disappeared before that time. At any rate there appears
to be no multiplication either in the sea-water or in the
body of the shell-fish—on the contrary, we found, as
others* have done, that in clean sea-water the Bacillus
typhosus rapidly decreases in numbers.

In further experiments where the infected oysters
were subjected to a running stream of clean sea-water, the
results were definite and uniform. There was in all cases
a great diminution or total disappearance of the typhoid
organism in from one to seven days. The stream of water
enables the mollusc to purify its gills and alimentary
canal, and so free itself from the results of sewage
pollution; and we found that in the great majority of
cases most of the bacteria were in fact cleared out in the
course of the first three days.

A considerable amount of attention was also given
in these reports to other diseased conditions of the oyster,
and to the presence of copper and iron in abnormal
quantities in the tissues of shell-fish from some localities.

Some of the earlier Lancashire Sea-Fisheries
Laboratory Reports, from 1895 onwards, gave brief notes
of the work that we were doing on the bacteriology of the
oyster, covering much the same ground as the reports that
were made to the British Association; and as a general
Summary, in the Lancashire Report for 1903 I had an
article, entitled ‘‘ Sewage and Shell Fish,’’? which dis-
cussed the evidence that had been accumulated locally

* De Giaxa has shown that even if pathogenic bacteria are able to
live for a time in sterilised sea-water they soon die off in the struggle
for existence with the bacteria of normal sea-water.

166 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

(and which had been laid before the Royal Commission on
Sewage Disposal* then sitting) in regard to the serious
contamination by sewage of some of the shell-fish beds
of the district. This article included the report by the
late Mr. R. A. Dawson, then the Superintendent of the
Fisheries District, on the mussel beds of our coasts in
regard to danger of pollution by sewage, revealing a very
serious state of affairs in some parts of the district.

In connection with this article in the 1903 Report we
have the first of Mr. Johnstone’s papers on the bacteriology
of samples of mussels from the Mersey Estuary. This
report was submitted to the Lancashire Sea Fisheries
Committee and was also communicated to the Board of
Agriculture and Fisheries at their request. Mr. John-
stone demonstrated the presence in these mussels of
bacilli which were regarded as affording certain evidence
of sewage pollution. This investigation was followed by
the examination of samples of mussels from various other
beds in the district; and in all these cases in which
sewage contamination was reported upon from this
Laboratory we did not rely upon the bacteriological
evidence alone—that evidence throughout has been used
as an important corroboration, but not as the sole proof.

In the fourth report of the Royal Commission on
Sewage Disposal, 1904, the Commissioners state that they
would not be justified in recommending that the closing
of a shell-fish bed or laying should depend as a matter of
routine on the results of the bacteriological examination,
and this is very much the conclusion at which Mr.
Johnstone and I had then arrived as the result of our
experience, and it is the opinion that I gave in my
evidence before the Royal Commission. In 1904 I
expressed my views on the question of samples as
follows :— ne gaa

* See Fourth Report of the Commissioners, Cd, 1884, p. 90, 1904,

SEA-FISHERIES LABORATORY. 167

‘In taking samples of suspected shell-fish I would
attach great importance to personal supervision by a
scientific or fisheries expert. The samples should
obviously not be taken by the parties interested, and they
should not be taken by disinterested, but untrained
collectors who may miss seeing some qualifying factor or
some important piece of evidence. A knowledge of the
local conditions, of the influence of tidal and other
currents, and of prevalent winds, may be of great value
in judging of the presence and extent of pollution, and of
the parts [of the bed] liable to be affected at a particular
time of day, or of the month. Consequently a personal
examination of the locality by a scientific man is always
important. Samples from various parts of the same bed
may have to be taken at different states of the tide, and
these should be chosen with knowledge and discrimination.

“Any additional evidence that can be obtained from an
inspection of the physical and biological conditions on the
bed is all the more important because of our want of exact
knowledge as to the meaning and value of some bacterio-
logical results. The topographical observations and the
laboratory work ought always to be considered together,
and must be regarded as parts of the same investigation
conducted by the one Authority. The bacteriological
examination may at once confirm the field-work in such
a manner as to leave no doubt as to the purity or pollution
of the locality, or it may give useful indications which
suggest the necessity for further observation of the local
conditions. It may also give a measure of the amount of
pollution. The question has been raised as to whether it
is possible to fix a standard of pollution which should be
regarded as dangerous to health. Can we say that all
samples yielding say 10, or say 20 B. coli per c.c. must be
condemned, but that those showing less than say 5 per

168 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCTETY.

c.c. may be tolerated? Before answering such a question
we must have further investigations. There are still too
many of the points involved which are left in doubt. For
example, we cannot be certain that all samples yielding
10 B. colt per c.c. are equally dangerous. Even if we
assume (as we probably may safely do) that pure oceanic
sea-water is free from B. coli and allied organisms, and
that these are to be taken as an indication of some sewage
contamination, we do not know how remote in time the
pollution may have been and how comparatively harmless
from a pathogenic point of view it may have become. It
is possible, or even probable, that B. colt may be
distributed to considerable distances in the excreta of fish
and sea-birds, possibly with some modification. Then
again, the bacteriology of the shrimp’s alimentary canal
requires examination, and we may add the fishes that feed
upon the shrimp. There are also other sewage feeding
invertebrates that may conceivably pass on some organisms
and not others, and may favour the distribution of B. colz
under circumstances that deprive its presence of any
special [pathogenic] significance.

‘“T am not arguing against the value of bacteriology,
but against a possible abuse of the method, and in favour
of a much wider investigation in which the laboratory
work will in all cases be supplemented, guided and
inspired by the marine biologist’s work in the field. The
case of each estuary, bed, or laying must be regarded as
a separate problem to be solved with a full knowledge of
all the local conditions.”’

In 1905 Mr. Johnstone reported to our Scientific
Sub-Committee on the detailed examination of the Llan-
fairfechan mussel bed in the Menai Straits. He and Dr.
Jenkins personally collected samples for analysis and
studied carefully the topographical relations of the shell-

ie hee pe, ee a

SEA-FISHERIES LABORATORY. — - 169

fish on the bed in relation to sewage outflows and dispersal
and defects in the drainage arrangements. As a result of
the examination it was found that Bacillus coli could be
isolated from the majority of the mussels examined. The
presence of this microbe usually indicates the contamina-
tion by faecal matters of the shell-fish in which it is
found. But unhappily, this organism must now be
regarded as present almost everywhere in shell-fish
bedded on our coasts, and its significance lies not so much
in its mere presence as in its relative abundance. It will
be seen from the detailed results stated in the Report
that B. coli was very abundant in several of the mussels
examined: these were the ones collected from the piles
in the neighbourhood of a break in the pipe. Two
mussels in the sample were quite sterile, and in one
or two others the microbe was present in very small
quantity; these latter shell-fish were collected from the
piles some considerable distance from the break in the
pipe. Only in one or two of the mussels examined was
the degree of pollution at all excessive.

In this case Mr. Johnstone points out that
“it is probable that the faulty condition of the sewer
pipe is the cause of the greater part of the pollution of
these mussels. The eddies caused by the tide round the
piles have excavated a shallow gutter directly beneath the
sewer pipe. As the tide lays bare the sands, this gutter
becomes filled with a mixture of sea-water and sewage
flowing from the break in the pipe. Then when the tide
begins to flow, some of this water becomes washed up
against the mussels on the piles, and the former become
polluted. If the sewer pipe were in proper repair, so that
all the discharge flowed from its extremity, and still more
if there were an intercepting tank at the pumping station
and the sewage were only liberated on the ebb tide, the

170 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

pollution of the mussels would be very slight, and would
be due only to the slight amount of general pollution of
the sea caused by the sewage from the towns at the
entrance to the Menai Straits.’

These conclusions, at which Mr. Johnstone arrived
from an examination of the conditions on the ground
quite as much as from the bacteriological investigation in
the laboratory, illustrate the value of a personal topo-
graphical inspection by a competent marine biologist.

In continuation of the experiments which I had
begun with Prof. Boyce, on cleansing shell-fish from
sewage bacteria by means of currents of water, I asked Mr.
Andrew Scott, in 1904, to re-investigate the matter with
polluted mussels at the Piel laboratory in the Barrow
Channel. Mr. Scott used mussels taken from a sewage
polluted area, and after ascertaining that they contained
large numbers of sewage bacteria, he exposed them to
currents of 1°75, 3°3 and 6 gallons of clean sea-water per
hour. The results of a number of experiments showed a
rapid diminution in the number of bacteria as measured
by the colonies produced in Petri dishes of neutral-red
bile salt agar. At the commencement of the experiment
the control mussels showed numbers like 1200, 1500 and
2000 colonies, and at the end of from 24 to 48 hours
numbers such as 30, 20, and 10 colonies only were found.

In illustration of the importance of understanding
the tidal and other currents before collecting samples of
water for bacteriological examination, take the following
series of observations made by Mr. Johnstone. In May,
1908, he made comparative cultures of samples of water
taken from the Barrow Channel every two hours, and
obtained the results given in the following table, stated in
number of intestinal bacteria in 2 c.c. of water :—

SEA-FISHERIES LABORATORY. LiL

FLoop TIDE WATER Ess TIDE WATER
5 hours before high water ... 0 64 hours before low water ... 0
ee 9 %5 eel ee 2 me wae oO
wv :,, 2 5 ssenO) ae Ks u «oa, 200
eet. » » ..-L000+

These investigations and results, which might be
extended with advantage to other channels and estuaries,
are of considerable importance. They indicate that even
in an estuary which was proved to be polluted the flood
tide water may be clean. A practical application of this
fact is the possibility of cleansing polluted shell-fish in
the flood tide water intercepted in shore ponds or tanks.

The numbers of organisms in 1 c.c. of water may
vary also on different parts of a shell-fish bed as the result
of amount of shelter, exposure to prevalent winds and
many other local factors which must be studied on the
ground by an expert and which make every shell-fish bed
an independent problem to the marine biologist. Con-
sequently, it is sometimes possible to collect samples of
relatively pure and of highly polluted mussels from
neighbouring parts of the same bed. Mr. Johnstone has
dealt with these considerations in some detail in a recent
paper (Journ. of Hygiene, February, 1910) from which I
quote the following paragraph :—

‘It is clear that much may depend on the precise
conditions under which the sample is taken, and upon
the precise spot. If this is so, then great caution is
necessary in applying the results of analysis of a sample.
of shell-fish purchased from a market stall or shop, to the
general locality from which the molluscs are said to have
been taken. Not only so but the Report must, in justice
to the fisherman, consider the length of time which has
elapsed since the shell-fish were taken from the sea, and
the conditions under which they have been stored. ....

172 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

What then is to be said of the interpretation of the
results of the analysis of shell-fish which may have been
taken from the sea some six days before the date of the
sampling, and which may have been stored in insanitary
conditions in the meantime? The discovery of pathogenic
organisms in such shell-fish might indeed be conclusive
proof of the origin of a disease or epidemic, but the
tracing of the latter to the part of the sea from which the
shell-fish were alleged to have come might be erroneous.
It is surely unfair to condemn a locality on the results of
such an analysis made perhaps on moribund animals in
which partial decomposition may have begun.”

After the first few years of work (1895-99) in which
Sir Rubert Boyce had collaborated, the Lancashire Sea-
Fisheries investigations were left entirely to the Fisheries

Laboratory in the Zoological Department, and _ the

bacteriology of the shell-fish beds from this time onward
has been in the hands of Mr. James Johnstone, B.Sc.,
under my general direction—and the practical work, both
on the shore and in the laboratory, has always been
carried out by Mr. Johnstone or under his personal super-
vision. Most of the Annual Reports on our Scientific
Fisheries investigations for the last ten years contain
papers on the bacteriology of the local shell-fish beds by
Mr. Johnstone, and in addition he has carried out other
inspections of suspected localities which led to private
reports to the Committee not published in our Annual

~ volume. All the “beds” and “layings” in the district

have been surveyed and charted, every sewer outfall has
been examined with respect to its influence on adjacent
shell-fish, and systematic bacteriological analysis has been
carried on every year. This constitutes a considerable
mass of scientific work which has caused the Lancashire
shell-fish beds to become better known bacteriologically

SEA-FISHERIES LABORATORY. 173

than those, probably, of any other part of the coast. I
shall only refer now to a few of Mr. Johnstone’s investi-
gations as examples : —

In 1904, Mr. Johnstone reported on the mussel beds in
the Mersey Estuary and in the Estuary of the Lune, and
on the deep-sea oysters of our district. In this paper the
dangerous pollution of the Egremont mussels was clearly
demonstrated, amounting in some cases to about 300
colon bacilli in a drop—say one-tenth of a cubic centi-
meter—of the stomach contents of the shell-fish. The
Rock Ferry mussel bed was also shown to be grossly
polluted in some parts. The Wallasey bed, although
yielding some evidence of sewage, was clearly much less
polluted than in the case of the Egremont and Rock Ferry
mussels.

The examination of the Lune Estuary was made at the
request of the County Council; and Professor Delepine,
of Manchester, also made an independent investigation
and report. The results showed the presence of sewage
organisms, but that compared with the Egremont mussels
the pollution of the Lune mussel beds is not excessive.
Finally, “ deep-sea ” oysters from various off-shore fishing
grounds were found to be unpolluted.

In this paper Mr. Johnstone doubts the practi-
cability of erecting a permissible standard of impurity.
The deep-sea oysters would satisfy any standard, the
Egremont beds would be condemned on any adequate
standard, but the Wallasey beds are in an intermediate
condition, and there might be difference of opinion
amongst experts as to whether the standard shouid pass
or condemn that degree of pollution.

In the Summer of 1906, Mr. Johnstone furnished a
report to our Scientific Sub-Committee on the state of
affairs at St. Annes-on-the-Sea, in which he showed that

174 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

this was a locality where the grave pollution of the
mussel beds is due to the direction of the tidal streams.
He says, ‘‘ The topography of the coast, indeed, renders
it impossible that these shell-fish can escape direct
contamination.’’ The actual pathogenic organism of
enteric fever, B. typhosus, was isolated from the mussels
examined from this bed.

In the same year (see our Annual Report for 1906)
Mr. Johnstone commenced an extensive investigation of
the mussel beds at the “ Ring-Hole,” and elsewhere in
the neighbourhood of Morecambe, a matter of great
importance on account of the various interests involved—
including the transplantation of ill-nourished, stunted
mussels from other localities to this more favoured
spot for active growth. ‘Ihe bacteriological results
showed that Bacillus colt was present in practically
all the mussels examined, but Mr. Johnstone urges
that to describe a mussel as dangerously polluted
merely because it contains Bacillus coli would be quite
unjustifiable. Its presence only indicates the possibility
that the shell-fish in which it is found may, under certain
circumstances, harbour strictly pathogenic organisms such
as the typhoid bacillus. When the number of Bacillus
colt in a mussel is small, then this possibility is remote.
When the number is very large, or when the pollution is
notorious (as in such cases as those of the mussels at
Egremont, Rock Ferry, or St. Annes), then we may
reasonably conclude that the shell-fish should not be
used for human food. We agree with the Sewage
Commissioners when they say, “If it should be seriously
contended that the mere presence of Bacillus coli or coli-
like microbes in an oyster should condemn it, few oysters
would probably escape condemnation.” The same is no
doubt true of mussels.

| ee

SEA-FISHERIES LABORATORY. ee

Of course it would be desirable, were it possible,
that no crude sewage, nor even purified effluent, should
find its way to any shell-fish bed ; but when one considers
the dense population of the littoral of Lancashire, it is
evident that, with our present knowledge and appliances,
the ideal of removing all sewage, so that no trace of it or
of its organisms can reach the shell-fish beds, is quite
unattainable.

Hitherto our bacteriological work had been mainly
on Oysters and Mussels, but about this time, 1907-8, Mr.
Johnstone commenced a systematic examination of the
Cockle beds on the Lancashire littoral, and made various
inspections, analyses and reports (see our Annual Report
for 1908). The methods of observation and investigation
are described in detail in order that future examinations
of these and other beds in the district may be conducted
on the same lines, so that the results may be comparable.

As a result, the numbers of the “coli” group of
organisms per cockle, in the six chief beds examined,
were found to be as follows :—

Ansdell ele O15 Silverdale ee 28
Formby hh ama OHO). Flookburgh __... Li
Leasowe_.... 60 Southport ay 12

The cockles from Ansdell are seriously polluted.
Formby is a doubtful case, perhaps Leasowe is also ;
but the others are so slightly affected that probably the
few organisms present represent merely the general dis-
tribution of “‘coli’’ in our coastal waters, and have no
harmful significance. A thorough investigation of the
distribution of the colon bacillus in the coastal waters
of the Irish Sea would be of great interest and practical
value.

About this time, the late Dr, H, Timbrell Bulstrode,

176 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of the Local Government Board, who had been interested
in our earlier observations on oysters in connection with
disease, now began to come into close relation with our
work on mussels and cockles. He took part with Mr.
Johnstone in visits of inspection to beds in Lancashire and
North Wales, and came to Liverpool on several occasions
to consult with usin regard to the co-relation of bacterio-
logical and topographical evidence and the possibilities
of remedial measures—a matter in which he took the
keenest interest. Dr. Bulstrode’s premature death iast
year is a severe loss to public health sanitation in con-
nection with shell-fish, a subject he had made peculiarly
his own by the two admirable Reports he prepared for
the Local Government Board, the first on oysters, pub-
lished in 1896, and the second on other shell-fish in 1911.

We have been in communication also in recent years
with other Government Officials, and with Public Health
Officers of several neighbouring towns and districts, and
some of our recent bacteriological investigations were
undertaken in consequence of these communications. It
was, for example, a statement received from the Medical
Officer of Health of an inland town that gave rise to
Mr. Johnstone’s report on the bacteriological condition
of the mussel beds in the Estuary of the Wyre. It is
clear that although the degree of pollution is not so great
there as in other known cases, the examination was very
desirable, and further investigation of that neighbourhood
may become necessary. The importance of such work,
carried out in co-operation with the Public Health Officers
of the towns to which shell-fish are consigned, can
scarcely be over-estimated, and it is gratifying to find
that Dr. Bulstrode, in his recently published “ Report
on Shell-fish other than Oysters in relation to Disease”
(Local Government Board, Cd. 5313, 1911), refers in

SEA-FISHERIES LABORATORY. Way!

appreciative terms to the investigations on shell-fish, both
under healthy and unhealthy conditions, which have been
carried out by the Lancashire Sea-Fisheries Committee.

Dr. Bulstrode’s companionship and co-operation in
these investigations have been of great value to us, and
it has been encouraging to find that we were generally
taking the same view of the problems that he did, and
were reporting on the various localities in very similar
terms. As a final reference to his recent Report, I may
point out, for the information of our local Committee,
that, in discussing the possible machinery for the
regulation of shell-fish areas, Dr. Bulstrode refers to the
further valuable work that might be carried out by a
Sea-Fisheries Committee organised for scientific research
as ours is, and says: “ The Officers of the Lancashire and
Western Sea-Fisheries Committee have, under the
guidance of the Honorary Director, Professor Herdman,
F.R.S., together with Dr. J. Travis Jenkins, Mr. James
Johnstone, and Mr. Andrew Scott, made a detailed survey
of the shell-fish beds and areas in their district, and of
the sewers and drains which discharge in their vicinity.
They have also from time to time made valuable
bacteriological examinations of the waters and shell-fish
in different places, and have conducted experiments as
regards the re-laying of shell-fish, which are likely to
prove of permanent value both to the shell-fish industry
itself and to the public health ” (loc. cit., p.. 123).

This recognition of the value of the Committee's
work in the Official Report of the Government Depart-
ment directly concerned should encourage and stimulate
those who are doing the work, and may justify the
Committee in undertaking a larger expenditure on these
very necessary investigations.

Turning now to the last, and perhaps the most

178 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

important piece of shell-fish work we have undertaken
in connection with public health, it was as far back as
the summer of 1906 that Mr. Johnstone, in consequence
of information we had received as to the condition of shell-
fish sent to the markets, commenced the investigation
of the mussel beds in the Estuary of the Conway, in
North Wales, an investigation which has been carried
on intermittently until the present time, and is probably
not yet finished. The Conway mussel industry is of
considerable importance. Amounts of up to 6,000 cwt.
per month (of the value of over £700) were sent from
Conway during part of 1906, to Manchester, Leeds,
Huddersfield, Halifax, Nottingham, and other inland
towns.

Sewage, however, is discharged into the Hstuary at
Conway, and also into the river above, and there
can be no doubt that a considerable degree of
pollution of the mussels is present. Several out-
breaks of enteric fever in inland towns have
now been attributed, by Public Health Officers,
to Conway mussels. Public enquiries have been held, and
a great deal of further work, both topographical and ~
bacteriological, has been done by Mr. Johnstone since
his preliminary paper in our Report for 1906. Moreover,
an examination of the beds was made along with Dr.
Bulstrode in 1907, and along with an Inspector from the
Fishmongers’ Company in 1908, and our results were
entirely confirmed by these independent authorities. But
although the Conway mussels are undoubtedly polluted
with sewage, stillit must be remembered that, in all such
cases, it is an unscientific and inconclusive statement te
attribute enteric fever in (say) Manchester to the mussels
of (say) the Conway Estuary, merely because such mussels
when eaten by the patient, or when purchased in the

SEA-FISHERIES LABORATORY. 179

market, were found to be sewage-polluted and to give
rise to the disease. In order to complete the case it is
necessary to exclude all other possibilities of contamina-
tion of the shell-fish between the date of gathering from
the mussel bed and that of sale and consumption in the
town. It is well known that shell-fish are sometimes
kept by the dealers for a considerable time before being
sold, and the possibility of contamination from sources
other than the natural habitat of the shell-fish must often
be a very real one.

As the Conway mussels were getting a bad name in
some markets, and as the industry was an important one
locally and well worth attempting to save, the idea*
occurred to us of trying to induce the fishermen to
establish a practice of transplanting their mussels for
a few days into cleaner water before sending them to
market. So with the object of having more definite data
on which to base recommendations, Mr. Johnstone made
a series of experiments during 1908 to ascertain the
conditions under which polluted Conway mussels would
cleanse themselves from sewage bacteria when placed in
unpolluted water in the Conway neighbourhood. A
locality at the entrance to the estuary, on the Morfa
beach, was selected because the water there was found,
as the result of detailed bacteriological examination, to
be much more pure than that of the Conway Estuary in
general. Several lots of polluted mussels were then taken
from the worst parts of the Conway beds and were placed
in boxes fastened to the shore at the chosen part of
Morfa beach. Samples of these mussels were examined
bacteriologically at the beginning of the experiment, and
further samples were examined 4, 8 and 16 days after

* The suggestion was also made by Dr. Klein in his Report to the
Fishmongers’ Company.

180 TRANSACTIONS LIVERPOOL BIOLOGICAL soOctETyY.

re-laying. The change after 4 days was most definite.
In the mussels as laid down the average number of
colon-like bacteria on each of the culture plates was
about 200; while in the case of those that had been
transplanted for 4 days the number in the similar cultures
was about 40. In most cases the reduction showed a loss
of 93 per cent. in the 4 days. In some cases the reduction
amounted to a total disappearance of the “coli”
organisms. It is clear then from these experiments made,
not in the bacteriological laboratory, but in the open
under natural conditions, that a very considerable degree
of cleansing follows when badly polluted mussels are
re-laid in unpolluted water, and that a period of four
days is long enough for practical purposes. After the
fourth day lttle or no further cleansing takes place.

The recommendation* was therefore made that
mussels taken from certain specified grounds which were
known to be badly polluted ought to be re-laid on the
Morfa beach for at least 4 days before being sent to
market. Compared with the capital value represented by
the Conway mussel ground, and with the increased sales
which would no doubt result from the confidence on the
part of the purchaser that the mussels were free from
danger, the initial and current expenses of this system
of re-laying at Morfa would be quite trifling. It is
surprising that the fishermen do not endeavour to make
their industry more lucrative by themselves putting
the recommendation into force.

The Corporation of Conway made an attempt last
winter to compel the fishermen to reform the fishery.

* The results of the experimental work were yéported to our
Scientific Sub-Committee and were discussed in February, 1909, and
again the following November; but it was found that the Sea
Fisheries Committee had no power to establish such cleansing depdots,
nor to make regulations enforcing their use if established.

SEA-FISHERIES LABORATORY. ~~ °° ~=—- 181

Impelled by the decline of the fishery, by the condemna-
tion of Public Health Authorities, and by our own adverse
reports, they applied to the Board of Agriculture and
Fisheries for an Order enabling them to improve, main-
tain and regulate the mussel fishery in the estuary, and
a Public Enquiry was held at Conway in December by
an Official of the Board. The object of the Order was to
enable the Corporation to take such steps as might be
necessary for the continuance and development of the
fishery, consistent with the protection of the public
health; and the provisions of the Order included the
establishment of cleansing depéts such as had been
recommended in Mr. Johnstone’s Report.

At the Enquiry the fishermen took up an impossible
position. They did not want the Order in any form,
they objected to the regulations suggested, and especially
to the proposed royalty of 6d. per cwt. to pay the expenses
of the improvements. They denied that the mussels
communicated disease, and asserted that there was more
demand for them all over England and Wales and
in Conway than there ever had been, and that they never
had complaints. They expressed the belief that ruin
would fall upon them and Conway if the Order became
law. The only object of the Corporation, in promoting
the Order, they said, was to get money. If it were con-
firmed ‘‘everybody would haul up their boats and
tackle.’’ In cross-examination they denied that they
washed their mussels near the sewers—a practice which
had been observed by both the late Dr. Bulstrode and
Mr. Johnstone when making their inspections, on several
occasions. This uncompromising hostility to regulation
on the part of the men must do harm to the industry.
The Conway mussels are regarded with suspicion, and it is
probable that the fishery, in the absence of any regula-

182 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,

tion, may still further decline; and in view of the
increasing pollution of the estuary, it may become a
dangerous source of epidemic disease.

As Mr. Johnstone has pointed out in his last report
to our Committee, this is all very regrettable, for there
is absolutely no doubt the fishery can be developed and
that risk of the communication of disease can be largely
if not entirely averted. All that is required for this
desirable consummation is the co-operation of the
fishermen with the Corporation, and the sympathetic and
intelligent regulation of the industry. There are, in my
opinion, no obstacles which will not yield to careful
study of the local conditions. Details of the construction
of cleansing ponds and storage of the mussels are not
likely to offer lasting difficulties. The mussels gathered
might be classed and the royalty charged in proportion
to their quality. When a man placed his shell-fish in
the cleansing ponds he might be given a receipt, on
production of which he would receive the same quantity
of cleansed mussels, of the same class, at the expiration
of a given time. The shell-fish would be sent to the
markets accompanied by a certificate, and public con-
fidence in their harmlessness would again be restored.

In his Report to the Local Government Board the
late Dr. Bulstrode saw no alternative, if this mussel
industry were to continue, but to close the major portion
of the Conway, and only to allow fishing on the seaward
parts of the estuary “as a provisional measure.” Hither
these measures are likely to be adopted by some Public
Authority to be created by legislation, or the Conway
mussels will gradually be excluded from the markets
under the pressure of the Medical Officers of Health in the
towns to which the mussels are consigned.

As an example of the situation hkely to arise in

SEA-FISHERIES LABORATORY. 183

connection with many of our shell-fish beds, we may
take the recent action of the Health Committee of
Birmingham in refusing to admit mussels from
a certain locality in Wales without a certificate of
purity. As a result the fishermen have applied
to our Committee to provide them with such a
certificate. This is a matter requiring careful
consideration. There is no doubt that such a certificate
ig wanted, and ought, under proper regulation, to be of
great service in the interests both of the public health
and of the fishing industries. But, if decided on, it must
be granted not in relation to one locality only, but for
each fishery of the District where the conditions are
favourable, and could only be given after adequate
inspection, and subject to periodic renewal. Unfor-
tunately it could not be given for certain localities
under their present conditions. Conditions, however,
keep changing. Both shell-fish beds and sewage
distribution undergo alteration, and so the pollution of
a locality may improve or may become worse. Not only
is each bed or laying a problem in itself, but it may be
a different problem next year from what it is now.
Moreover, in the case of beds that are condemned as
dangerously polluted, remedial measures might well be
undertaken. If compulsory cleansing of the living
mussels in tanks of purer water for a short period before
sending to market is not found efficacious, Dr. Bulstrode’s
suggestion of complete sterilisation by steaming, as is
the custom with cockles at Leigh, may have to be adopted.

It is evident that if the shell-fish from specified
localities in the District are or are not to be certified as
fit for human food, some standard of permissible
bacteriological impurity must be adopted. In respect of
shell-fish examination as a matter of official routine, this

184 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

question now arises for the first time in this county.
Hitherto no recognised standards have been established,
since the kind of work done so far has consisted of
investigation of the condition of shell-fish or shell-fish
layings with reference to some special question—an
epidemic, etc.—arising in relation to some specified
locality. If the Committee should decide to issue
certificates of purity—or withhold them—some standard
of bacteriological impurity, as well as some standard of
other available evidence, must be adopted.

It is fortunate that there are, in the Lancashire and
Western Sea Fisheries District, some shell-fish areas
which, so far as we can judge, are ‘almost free from
pollution—at least free from dangerous pollution.
Bacteriological results obtained from such localities may
assist in the formation of a standard. At all events it
will be necessary for the Committee, in the absence of
the guidance or the experience of other public bodies, to
set up its own criteria of what constitutes a dangerously
polluted mussel or cockle. |

It is clear, then, that a very important part of the
work of Sea Fisheries Authorities in the future, if our
great shell-fish industries are to be maintained, will be
the periodic inspection of the whole coast by competent
scientific men working on rational lines, such as those
adopted by the late Dr. Bulstrode, where all the factors
of the problem are taken into consideration. The
granting of a certificate implies the institution of some
standard of purity and in fixing this topographical details
must be fully considered ; and the results of the further
bacteriological analyses are to be interpreted in the
light of such details. It is doubtful whether we are
ever justified in applying the results of bacteriological

analysis alone in administrative routine. It would, no

SEA-FISHERIES LABORATORY 185

doubt, be very simple, and would seem desirable, if a
Local Authority were able to reject or approve a con-
signment of shell-fish on the results of a routine
examination in the bacteriological laboratory, and,

‘aif this were possible, much trouble would be

avoided. Unfortunately, this simple procedure is
not adequate, the trouble must be taken, .common-
sense topographical evidence must be considered
or in many cases unjustifiable hardship would
be inflicted on the fishermen, the industry -would be
seriously damaged, and the real source of pollution
might fail to be traced. Over and over again in our local
work we have come upon cases where either a favourable
or an unfavourable bacteriological report in regard to a
bed might have resulted according to the exact spot from
which samples were gathered or the precise conditions
under which they were taken. Moreover, in other cases
we have shown that the bacteriological results can only
be properly interpreted by those who have an intimate
knowledge of the natural history of the locality.

All this kind of work, in the interests both of the
public health and also of the fishermen who make their

living from the shell-fish industries, ought to be under-

taken in all cases by a biological Bacteriologist who is
at the same time a good Field Naturalist and a Fisheries
ixpert.

In a rational bacteriology upon which. regulation
of an industry may come to be based, and from which
conclusions as to the source and the history of the
infection may have to be drawn, it is not sufficient merely
to record the proportion of a certain sample of shell-fish
in which a certain organism was observed. For
example, the type of statement which one meets with
in the annual reports of Medical Officers of Health, that

186 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY

out of so many oysters or mussels examined during the
year a certain number contained ‘‘coli’’ and a certain

6€

number ‘‘ enteritidis,’’ 1s useless for all practical pur-
poses, and cannot lead to any result. As I have shown
above, the significance of ‘‘ coli’’ infection les not in the
mere presence but in the relative abundance of the
organism. Records of the bacteriology of samples of
shell-fish are futile unless one knows in detail where the
sheil-fish came from, the conditions under which they
were collected, their history since the time of collecting
and the relative number of each kind of organism present.
The presence of such an organism as B. colz in relatively
small quantity, and possibly of remote origin, may be of
no importance in connection with the public health, and
at most it merely indicates the possibility that the shell-
fish in question may under certain circumstances con-
tain pathogenic organisms. When the number of
organisms present is relatively great, and when there is
topographical evidence of comparatively recent con-
tamination, then the risk of pathogenic organisms
being present-—whether actually isolated or not—is
much greater, and condemnation of the shell-fish becomes
justifiable. But unless the relative amount of infection
is determined, and the various factors in the environment
affecting the problem are known in detail, the laboratory
bacteriologist or public health official runs some risk of
being deceived by the samples examined and of arriving
at erroneous conclusions, from routine analyses, as to the
real condition and history of the suspected shell-fish in
relation to sewage contamination.

In sea-fisheries investigation and administration we
must be careful that bacteriology remains our useful
servant and does not become a tyrannical master,

SEA-FISHERIES LABORATORY. 187

REPORT ON THE EXAMINATION OF THE
MUSSEL BEDS IN THE ESTUARY OF THE
Wee WITH: REFERENCE TO THEIR
HIABILITY ne CONTAMINATION in
SEWAGE.

By James JouHnstTone, B.Sc.

Karly in the present year (1911) Dr. Jenkins
received a communication from the Medical Officer of
Health of an inland town with reference to a consign-
ment of mussels said to have been sent from the Estuary
of the Wyre. A bacteriological examination of these
shell-fish had been made, and it was stated that the
analysis gave evidence of an undesirable degree of
pollution by sewage bacteria. In consequence of this
report Dr. Jenkins suggested to me that it would be
desirable to make an inspection of the condition of the
mussel beds in the Wyre area; all the more so since
those shell-fish have not been examined since the former
sewerage system of Fleetwood was replaced by that now
in operation. I accordingly made three visits to the
Wyre Estuary—on 9th February, 22nd February, and
9th March—and collected samples for examination on
the two latter occasions. I also received a further
sample of mussels on April 29th from Mr. John Wright,
who collected the shell-fish according to instructions.

I have much pleasure in acknowledging the
assistance and co-operation of Mr. T. R. Bailey, the
Port Sanitary Inspector, who accompanied me and
Mr. J. Wright on each of our visits of inspection. The
sketch chart of the sewage outfalls reproduced here has
been marked by Mr. Bailey from the Surveyor’s plans.

188 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY

The Fleetwood Sewage Outfalls.

The whole of the Fleetwood sewage was formerly
discharged into the Wyre at various points between the
docks and ferry, through separate outfall sewers, and it
is these that are shown on the chart published by the
Royal Commission on Sewage Disposal. At the present
time the untreated sewage of the Urban District (the
population of which in 1909 was estimated at 15,000)
is intercepted by a sewer which runs westwards towards
the Rossall shore, and discharges through an 18-inch
outfall sewer near Rossall Landmark. There is a storm
overflow near the pumping station on the Rossall side,
and another storm overflow discharges into Fleetwood
Harbour. There are two very small sewers near the
Knottend Ferry. The Wyre also receives the untreated
sewage from Poulton-le-Fylde from an outfall dis-
charging on the West side of the Estuary near Skippool
Marsh; and a small sewer discharges on the East side,
about half a mile above Wardley’ S Hotel near the brook
called Peg’s Pool.
| In addition to these outfalls there are drains
conveying effluents from (1) a fish oil works, (2) a fish
curing house, and (3) a fish meal works, all of which.
factories are situated on the side of the Estuary near
the docks. These drains are 6-inch pipes about 100
to about 300 yards long. They discharge on the beach
well above the level of low water of ordinary tides, and
the effluents reach the channel through little brooks, and
flow right over the mussel beds, between Preesall and the
docks. There is also a small drain: from an 1ce factory,
opening into the harbour.

It will be gathered from the foregoing dese nanan
and from the sketch chart, that the conditions in the
Wyre Estuary, as regards lability of sewage contamina-

SEA-FISHERIES LABORATORY. — 18y

tion of the mussels and oysters bedded there, are fairly
good. There can be no question of the Fleetwood sewage
fouling the mussel beds in the Wyre: it is discharged
on to the Rossall shore where there are no shell-fish beds
—at least none that need concern us here; and if we may
assume that the storm overflow in the harbour always
serves the purpose for which it was designed, we may

—EEEEEES

dismiss the question of the sewage of Fleetwood itself.
Two sources of pollution need only be considered,
(1) the effluents from the works mentioned above, and
(2) the sewage of Hambleton and Poulton-le-Fylde. It
was the presence of these sources of pollution that made

bacteriological analyses necessary.

190 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.
Bacteriological Analyses.

(1) Mussels from the beds near the docks (2 analyse).
(2) Mussels from the beds at Wardley’s.

(3) Estuarine water from the channel near the docks.
(4)

4) Water from the channel adjacent to the Wardley’s
Ferry Slip.

(0) Effluents from the fish oil and fish meal works.
Methods.

I give an outline of the methods of analysis;
obviously the comparative value of such analyses
depends on the methods employed. Grinbaum’s
neutral-red, bile-salt, lactose agar medium was employed
for the isolation of the bacteria regarded as significant
of faecal pollution.

Five mussels formed a sample. The soft parts were
removed from the shells and cut up finely with scissors,
then ground up in a mortar, and made up to a volume
of 250 c.c. with sterile water. One c.c. of this emulsion,
containing 1/50th part of a single mussel, was then
plated in the agar medium mentioned; five such plates
were made in each analysis.

One c.c. of the water was similarly plated.

In the case of the manufacturing effluents dilutions
were made so that 1 c.c. of each corresponded to 01,
0°01, 0°001 of the original liquid. 1 c.c. of the original
liquid, and 1 c.c. of each diluent, were plated as in the
case of the water. In all cases the usual precautions to
obtain sterility of the apparatus and materials used were
taken; and control plates to test the sterility (to the agar
medium employed) of the water used for dilution, the
air of the laboratory, and the hands of the operator,
were made. The results of the analyses were as follows:

(1) Mussels from the beds near the docks.—
(a) Taken at low water of a neap tide: The average

eC E—_

- =_—--

= a" oe
®

SEA-FISHERIES LABORATORY. 191

number of intestinal bacteria contained in one mussel
was 69. (b) Taken at low water of a spring tide: The
average number of intestinal bacteria contained in one
mussel was 90.

(2) Mussels raked from the channel near Wardley’s
Ferry Slip: The average number of intestinal bacteria
contained in one mussel was 150.

(3) Estuarine water from the channel adjacent: to
the place where samples (1) and (2) were taken: 1 c.c.
of the water contained, on the average, 07 bacteria
(less than one organism per 1 c.c.).

(4) Water from the channel adjacent to the
Wardley’s Ferry Shp: 1 c.c. contained, on the average,
22 intestinal bacteria.

Those unfamiliar with the bacteriology of shell-fish
and sewage effluents may best appreciate the meaning
of these results from the following comparisons. As a
standard result which may enable us to assign a value
to the Wyre analytical figures we may take the case of
the Kstuary of the Conway River in North Wales,
where the sources of sewage contamination are abundant
and obvious; and where there is actual epidemiological
evidence of the transmission of enteric fever by the
mussels taken from the estuary. Mussels taken from
these beds contained, on the average, about 2,000
intestinal bacteria each, the analysis being made by
methods identical with those described above.*

*Mussels bought from a retail shop may be very much worse
than those taken from a badly polluted Estuarine area. In one
sample bought from a low-class Liverpool fish-shop I found—as the
average of ten mussels—that each shell-fish contained 17,150
intestinal bacteria. It must often be the case that multiplication of
the contained bacteria—perhaps even direct re-infection—may take
place during storage in insanitary premises; and obviously it would
be unfair to lay the blame of such excessive pollution on the natural
conditions of the beds from which the shell-fish were collected.

= LOTTIE

192 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

In this case the water of the Estuary contained, at
the worst place sampled, 156 intestinal bacteria per c.c. ;
and, on the average, 35 per c.c.

Judged, then, by such standards the mussels taken
from the Wyre near Fleetwood Docks, and from the
channel at Wardley’s, do not appear to be dangerously
polluted; and the water in the lower reach of the
estuary 1s also passably clean. Two matters, however,
appear to require special consideration. These are :—

(1) The pollution of the Estuary at Wardley’s.

It will be seen that the average number of
intestinal bacteria contained in the surface water near
the Wardley’s Ferry is too high to be neglected. At
this point the influence of the pollution from Poulton,
and that of the outfall sewer at Peg’s Pool, are felt.
Nevertheless, the mussels taken from the bottom of the
channel at Wardley’s are very similar as regards
contamination to those from further down the Estuary.
The sample taken was practically fresh water, and the
sewage probably floats on the surface and may not
generally come in actual contact with the mussels on the
bottom of the channel. The mussels on the foreshore
near low water mark may, at times, give evidence of.
more serious pollution, but it does not appear that these
are taken for immediate marketable purposes. The
conditions at this part of the Estuary might be improved -
if the Poulton sewage were intercepted and discharged
for a short time only at the beginning of ebb tide. It
would then be greatly diluted and would not seriously
affect the shell-fish at the bottom of the channel. So
far as the results of these analyses go there does not
appear to be much danger of pollution by the Poulton
sewage, but nevertheless the relatively high bacterial

— es

i i ee a

—=— Tee

ee ae ee

SEA-FISHERIES LABORATORY. 1938

contents of the estuarine water at Wardley’s does
indicate the possibility of a fouling of the mussels under
certain circumstances.

(2) The Effluents from the Fish Refuse Works.

Cultures were made from the effluent proceeding
from the Fish Oil Works with the object of isolating
any intestinal bacteria that might be contained therein.
Plates inoculated with 1 c.c., 01 c.c. O01 c.c. and
0-001 c.c. were made, but all were sterile, Bacillus coli
being certainly absent from these quantities of the
effluent. _

Similar quantities of the effluent from the Fish
Meal Works were also analysed. Plates containing
01, 0:01 and 0001 c.c. were sterile after 48 hours’
incubation; but the plate containing 1 c.c. showed a
small patch of colonies. This, however, was, I think, due
to accidental contamination from a pipette used in
inoculation.

Ordinary domestic sewage may be taken as con-
taining from 10,000 to 1,000,000 Bacillus coli per c.c.
The effluents in question are no worse than the estuarine
water, and so far as this analysis goes they need not be
regarded as contributing to the pollution of the mussels
by intestinal bacteria. 7

These effluents are offensively smelling liquids. As
discharged on the beach they were, when I saw them,
clear, rather warm, and sometimes oily looking. The
smell was not that of putrefying organic matter, but
rather suggested aromatic compounds of some kind:
phenol, however, was not present in appreciable
quantity. Samples were placed in sterile flasks and
incubated at ordinary room temperature, but the liquids
did not contain any appreciable quantity of putrescible

194 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

substance, and the smell disappeared completely after
two days. They were very slightly turbid, but no more
so than ordinary river water.

In February, 1908, a sample of one of these
effluents was sent to me for examination. It was highly
septic, containing (in nutrient agar cultures) over
100,000 and less than 1,000,000 bacteria per c.c. It
contained no antiseptic substances, and when dosed with
sugar an abundant growth of bacteria and infusoria
appeared. It had the same offensive smell possessed by

the effluents examined this year. It was subjected to

chemical analysis at the County Laboratories, and gave
the following results :—
In parts per 100,000.

Total solid matter in solution... 2) Gs awZ
Oxygen required to oxidise in i .
(1) 15 minutes: ... Sa sh 37
(2) 4 hours Be ae Ga MES
Ammonia ae ay Ve Bis 12°5
Ammonia (from organic matter by dis-
tillation with alkaline permanganate) 54
Sulphuretted hydrogen — ins ike 0°36

Mixed with tap water (10 per cent. effluent and
-90 per cent. water) and allowed to stand three hours, the
mixture lost five-sixths of its total dissolved oxygen;
and allowed to stand twenty hours it lost twelve-
thirteenths of its dissolved oxygen. The Analyst
reported that “‘ Fish would be suffocated for want of air

in such a mixture of effluent and water, or even with a

smaller proportion of effluent.”

I did not collect this sample myself, and am
therefore unable to say whether or not the effluent may
have been accidentally contaminated by bacteria
during collection.

Fy -) —— op ——

— ————

— ———

———— a a eS

SEA-FISHERIES LABORATORY. 195

It does not appear probable that there is any danger
of the pollution of the mussels by human or animal
faecal matter by means of these effluents, and this is the
danger with which we are more immediately concerned.
But some attention should be directed to the pollution
of the foreshore and shell-fish by a discharge which
appears to be a noxious one, and which the Committee
may be empowered to prevent. Whether or not the
emanations from the Fish Oil and Fish Meal Works

constitute a technical “‘nuisance’’ is a matter for
consideration by the local sanitary authority; but the
fouling of the foreshore and shell-fish by an undoubted
manufacturing effluent (not a sewage effluent) is a matter
that comes within the purview of the local fisheries
authority. I would direct attention to the recently
published report on the shell-fish beds of England and
Wales by the late Dr. Bulstrode, of the Local Govern-
ment Board, in which this particular case of pollution 1s
considered.*

The effluents in question are described as offensive
ones, and it is stated that the poor quality of the mussels
on the foreshore is due to the detrimental effect of the
effluents. In the Analyst’s Report quoted above, the
opinion is expressed that the liquids would be harmful
to fish life because they would deprive the estuarine
water of a large part of its contained oxygen. It is true
that, as a rule, the discharge would be greatly diluted,
still it would probably happen repeatedly that small
parts of the mussel bed on the foreshore would be bathed
in effluent which had undergone little dilution—even
dilution to the extent of 90 per cent. would be likely to
be prejudicial according to the chemical analysis—and

* Supplement to Rept. Med. Off. Loc. Govt. Bd. for 1909-10—
Shell-fish other than oysters in relation to disease, By H, T. Bulstrode,
M.D., p. 222; [Cd. 5813] 1911,

196 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

to that extent the discharge would be noxious to the
fisheries.

I do not know what is the precise nature of the
manufacturing process at either of the works mentioned ;
or whether the effluents are always of approximately the
same nature. Obviously further examination, from the
‘point of view of the fisheries, is desirable,

SEA-FISHERIES LABORATORY. 197

SNe NStTVE STUDY OF THE MARINE
PLANKTON AROUND. THE SOUTH END OF
THE ISLE OF MAN.—PART V.

By W. A. Herpman, F.R.S., and Anprew Scort, A.L.S.

METHODS.

The work was carried on during 1911 on the same
lines as in previous years. Mr. W. Riddell again gave
most efficient help at sea, in the observations taken from
the yacht; Mr. Chadwick and Mr. T. N. Cregeen, of the
Port Erin Biological Station, collected the samples from
Port Erin Bay throughout the year; the two authors
divided the rest of the work as before; and Miss H. M.
Lewis, in the Zoological Department of the University
of Liverpool, devoted a great deal of time and trouble to
compiling the statistics, tables, curves and diagrams
from which this paper is written.

The work at sea from the steam-yacht *‘ Runa’’ was
carried on for some weeks in April, and again in the later
summer (August and September), usually the two most
important times of planktonic change. During the rest
of the year, statistical weekly gatherings were taken for
us in Port Erin Bay, in accordance with a uniform plan,
by the staff of the Biological Station.

We do not propose this year to make such a detailed
statement of the results as we have done for previous
years, but rather to give conclusions and comparisons,
and to pick out for remark any matters that seem new or
unusual. Consequently we would refer readers who are
interested in a fuller discussion of any points we have
already dealt with to the preceding four parts of this
work, (See Reports for 1907-1910.)

198 ‘TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

MaTERIAL AVAILABLE.

The collections made this year have amounted to
over 500—within the limited area off the Isle of Man to
which this “‘ Intensive Study’’ apples. This series
compares with those of former years, as follows : —

| At Sea, from Yacht. In Bay
Year. throughout Totals.

Spring. Autumn. Year.

1907 218 279 138 635
1908 156 242 sek, 555
1909 329 147 231+49 756
1910 107 249 296 652
EOVT 120 84 me 314 518
Totals ... 930 1,001 1,185 3,116

These make about 1000 in each of the three vertical
columns—Spring, Autumn and Bay—and from over 500
to over 750 for each of the five years in question.

The remarks made in the previous Reports about the
nets used and the methods adopted apply again; but for
the vertical hauls we have used almost wholly the Nansen
net, either open throughout the haul or closed after
traversing a certain zone. The other nets used were
‘coarse’? and ‘‘fine’’ (No. 20 silk) horizontal,
“‘funnel’’ net, ‘‘ Otter’? net, ‘*’ medium Nansen,”
‘“large Nansen,’’ and “‘Shear’’ net. All these nets
and the methods in which we use them have been
sufficiently described in the previous parts of this study.

PLANKTON OF PORT ERIN BAY IN 1911.

The plan adopted for the last few years in regard to
the plankton samples from Port Erin Bay has been to
take two horizontal (coarse and fine nets) and one vertical
haul twice each week throughout the year—thus giving

SEA-FISHERIES LABORATORY. 199

six samples in the week. The twelve months in 1911 are
represented as follows :—

meee eb PE) TET |) EV |. Vo} VE) VIL} VIET | TX | X | XE) XO

Gatherings | 27 | 18 | 30 | 31 | 27 | 31 | 27 | 23 27 | 22 | 24 | 27

Kach month is thus represented by a considerable
amount of material, the average per month being about
26 hauls. The lowest monthly records are 18 for
February and 22 for October, and the highest, 31 for
April and June. It has to be added that in April,
August and September additional material, obtained

from the “‘ Runa’’ outside the bay, 1s also available for
comparison.

Treating these records in the same way as in previous

years and comparing the results we find that :—

(1) The monthly averages for the horizontal nets
are not quite so large as those of last year. In
1910 the highest was 63 ¢.c; in 1911 it was
46 c.c.

(2) The maximum was reached a month later—in
May in place of April. If we analyse this
maximum into its three most important
constituents, we find that in 1911:—

The Diatom maximum was in May.
The Dinoflagellate maximum was in June.
The Copepod maximum was in July.

The vertical hauls at the mouth of the Bay, as on
previous occasions, agreed well in their evidence with
the much larger bulk of material obtained from the
horizontal nets. The maxima in the vertical hauls were
at the times stated above and the quantities were :—

In 1910 (April and May) about 4 c.c. per haui.

In 1911 (May and June) about 5 c.c. per haul.

fe)

200 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

It may be noted here that in 1911 the hours of
sunshine reached the monthly maximum in May, and
the total number was considerably greater than that of
1910. (See below under Sunshine.)

If now we neglect the vertical hauls—which are not
directly comparable with the horizontal—and treat each
pair of coarse and fine net samples taken at the same time
as forming together a single double haul, we get the
following result for the twelve months:—

Double Average Diatoms. Dinoflag- Copepoda. Copepod Copepod

1911. hauls. catch. ellates. juv. nauplii.
January ...... 9 3°6 52,064 3,511 . 3,314 188 1,553
February ... 6 4-0 60,528 2,508 2,579 222 2,537
Marchy: 3... 10 4-4 245,851 2,495 1,013 175 3,062
AN jel Wee apne eaoe 11 9-0 240,446 901 2,752 246 3,520
May Wee tei 9 46:0 24,201,900 26,230 lich! 2,039 15,240
SMe: “wets eas Ld 37:3 3,767,835 50,365 25,285 1,762 44,820
Duly! sraecewes 9 18-8 8,209 18,967 75,533 1,229 62,618
AUSUSG Ze... 8 11-4 1,998 1,510 61,351 2,426 35,058
September... 9 15:3 928,501 8,818 31,651 2,426 44,244
“October ...... 8 14-5 4,742,791 10,510 18,559 1,700 32,058
November ... 8 5:5 506,729 6,574 20,741 894 8,058
December ... 9 3-4 124,144 5,131 10,492 124 5,190

This table serves for comparison with those we have
published for 1909 in Part III, p. 212, and for 1910 in
Part IV, p. 199; and shows very clearly the average
monthly catches, forming a simple curve rising steadily
from 3°6 ¢.c. in January to 46 c.c. in May, and then
declining to 34 ¢.c. in December. It also shows the two
maxima in the Diatom column, the first in May with over
24 millions and the second in October with nearly 5
millions. Finally the succession of maxima, Diatoms in
May, Dinoflagellates in June, and Copepods in July, is
clearly seen. These maxima are all large ones, in the
ease of Diatoms and Copepoda much larger than in the
previous two years, and in the case of the Dinoflagellates
only exceeded by that of 1910.

If we add together Diatoms and Dinoflagellates in

Pla ee i i 8 Ni ee

SEA-FISHERIES LABORATORY. 201

May and June (the months when phytoplankton pre-
dominates), and compare the total with that for Copepoda
(both young and adult) in July and August (months of
the zooplankton maximum) the contrast is obvious.

Phytoplankton. Zooplankton.

May + June ...... 28,046,330 98,333
July + August ...... 30,684 238,215

These are, moreover, not the largest hauls in any of
the cases, but only monthly averages; and in the right-
hand column it is only the Copepod zooplankton that is
taken into account. Still the differences are quite
sufficient to show the changes in the nature of the pre-
dominant plankton in passing from the one period to the
other.

It is always necessary to analyse the total numbers
for the days, or nets, whenever a sudden change is seen,
in order to determine what has caused the change. For
example, in 1911, on April 21st the total catch in the
coarse net went up to 22°5 c.c. from an average of about
4:0 c.c. during the previous gatherings of the same net
that month. The gathering immediately before, on
April 18th, was 4°5 c.c. Now this great increase in bulk—
about five-fold—was not due to any increase in numbers
in any one of the more important groups, as the following

figures will show :—
Diatoms. Dinoflag- Copepoda. Copepod

ellates, nauplii.
700 bee 4-5¢.c. 238,000 3,000 7,651= . 7,000
PARHIL BE crasacne 22-5¢.c. 194,000 0 2,403 4,800

It is evident that these figures for Duiatoms,
Copepoda, etc., do not account for the rise in volume of
April 21st.

It is necessary, then, to examine the specific details,
when we find that an increase in the number of Medusae,
polychaet larvae, fish eggs, and a few other larger

202 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

organisms occurred on the 21st, and caused the increase in
the volume of the catch although not in the number of
organisms caught.

In the case, on the other hand, of a sudden rise early
the following month from 7 ¢c.c. on May 4th to 20°5 c.e.
on May 10th, the number of Diatoms rose from 115,450
to 2,268,750, which quite accounts for the increase in
volume of the catch, and this was in fact the beginning
of the vernal Diatom maximum, and the numbers of c.c.
went on in the next few days to 357, 42°8, etc., all caused
by Diatoms in tens of millions. As an example of how
thick the water was with phytoplankton. at the time, it
may be stated that on May 16th, when the largest’
horizontal hauls were obtained (42°8 in coarse, and 60°2
in the fine net), the vertical net hauled through only six
fathoms gave 17°2 c.c. (a very large haul for the vertical
net) and contained nearly three millions of Diatoms. As
a contrast to that, we have a month later the vertical net,
on June 15th, giving 16 c.c. (nearly the same volume of
catch) with only 8,200 Diatoms. On looking into the
specific details the cause is seen to be the great increase
in the number of Copepoda on the latter date.

Bay DIATOMS.

The following notes as to the occurrence of the
Diatoms at Port Erin at the time of the vernal maximum
were taken at the time of collecting, but have been revised
and added to since as a result of examining the catches in
~ detail :—

May 13th.—The Vernal Diatoms now appeared in
quantities (calm weather with a marked rise in
temperature).

May 16th.—Tow-net gatherings large, and consisted
almost entirely of Diatoms (weather continues

ee i

SEA-FISHERIES LABORATORY. 203

calm and the increase in temperature is main-
tained).

May 19th.—Diatoms occurred in very large quan-
tities, especially in the fine net.

May 22nd.—Catches rather smaller, but Diatoms
still in abundance, even in the vertical net
(weather still fine and warm).

May 25th.—Diatoms much less numerous. Fine net
had only about one-tenth, or less, of the
gatherings on May 19th (no obvious change in
weather conditions).

All the above large catches of Diatoms consisted
almost entirely of species of Chaetoceras. It was not
until a week later that Rhizosolena (chiefly R. semi-
spina) made its appearance. It reached its maximum
early in June, and then gradually died off. By the
beginning of July the Duiatoms had practically
disappeared.

The following gives the quantity of plankton and the
total number of Diatoms present in each haul of the fine
net taken during the month of the Diatom maximum : —

Date. Quantity ine.c. Total Diatoms.
Rita 1.4,35)..b>2.. 2-5 43,360
‘a eS ee 1-0 10,610
_- 0a Soe 6-5 525,680
A 30-2 19,118,000
~~ Ce 60-2 54,141,500
ETE S citesip ows ne 54:5 34,447,500
SRA Dengcinaenanes 30-5 27,775,000
aS 8-3 2,504,500
a> 14:8 22,023,100 ©
RS LS pcccw cance - 11-3 4,926,000
Be eas cet ened 24-7 12,943,000
- Sea aves « 12:7 2,656,000

The sudden rise on June 3rd is due to Rhizosolenia
semispina. The figures given above for May are
unusually large, and the increase from about ten
thousand on May 4th to over fifty-four millions on May

204 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

16th is most rapid. The most abundant species were
Chaetoceras debile and Ch. sociale. On May 16th the
first of these species reached 30,000,000, and the second
12,000,000 per haul.

THe More Important GENERA OF DIATOMS.

We have taken out again the same seven genera
as in our last report, viz., Biddulphia, Chaetoceras,
Coscinodiscus,° Rhizosolema, Thalassiosira, Guinardia
and Lauderia. ;

We think it unnecessary to print the detailed tables
again this year. Those we gave last year (Part IV, pp.
204-6) form a very good example, and we shall merely
note here the points in which the details for each of the
seven genera differ in 1911 from those recorded for 1910.

Biddulphia.—Does not attain to quite such high

numbers in spring this year, and the highest record
(312,560) 1s in April in place of March; but after dying
out completely in summer, the genus has a second, and
this year a greater, maximum in November (660,600 on
November 24th), when there is the high monthly average
of 341,231—last year it was under 40,000.
_ Chaetoceras.—Last year the maximum of this genus
(49 millions) was in April. This year it is a month later
with 68 millions on May 16th. The autumnal, smaller
maximum is larger than in the previous year, and reaches
over 10 millions on October 2nd, and about 6 millions on
October 16th and 19th.

Coscinodiscus.—The spring maximum was earlier
this year than in 1910, and reached 392,400 on March
14th. An unusually late haul was 79,290 on May Ist.
The autumn increase was unusually great, the maxima

being 42,400 on October 10th, 44,700 on October 16th,

SEA-FISHERIES LABORATORY. 205

and 40,450 on November 27th, as against 11,400, the
highest number in these months in 1910.

Rhizosolenia.—This important genus again had its
maximum in June, but did not reach so high a point as
in 1910. Notable hauls are 2,160,000 on May 22nd,
2,880,000 on May 29th, four hauls of from 34 to over
10 millions between June 3rd and 8th. The autumnal
increase this year did not amount to much, the largest
haul being 276,000 on October 19th.

Thalassiosira.—The spring maximum was again in
May, with 1,120,000 on May 29th. Quantities of several
hundred thousand per haul remained until June 5th, and
then the genus suddenly disappeared and was unrepre-
sented until October 5th, when 287,000 were taken in one
haul.

Guinardia.—This genus is very poorly represented
this year, the highest figure being 204,000 on June Ist,
compared with nearly nine millions the previous year.

Lauderia.—This form also shows smaller numbers
than in 1910. The only haul of over a million was
1,203,500 on May 19th—whereas it attained to 20
millions in April, 1910.

We add here the monthly averages of these seven
genera of Diatoms, as follows :—

1911. Biddul- Chaeto- Coscino- Rhizoso- Thalassi- Guinardia. Lauderia.

phia. ceras. discus. lenia. osira.
fibige 31,758 11,903 5,119 64 0 22 0
i aaaes's 38,150 8,606 11,572 50 Ona. 133 33
arch....... 124,225 41,990 73,796 1,425 0 283 160
ae 115,823 86,185 34,890 1,650 0 82 150
as od 21,139 22,745,683 17,531 858,208 217,329 27,938 312,363
aa a 2,073 1,431,005 200 2,121,226 135,073 60,656 12,864
ae 0 478 0 7,180 0 551 0
219 1,437 12 310 0 31 0
ae 11,110 868,017 1,479 21,018 0 481 1,511
Ric's’ 142,690 3,956,047 25,444 72,622 84,462 987 41,387
ies 341,231 143,251 17,402 0 12 2,597 12

seneee 77,788 31,159 9,411 0 0 4 0

206 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The above table brings out clearly the marked
Diatom maximum in May, the minimum in July and
August when very few Diatoms were present, and the
second or autumnal maximum in October. The vernal
and autumnal maxima are shown unusually clearly by
'halassiosira, which was present only at these periods,
being totally absent from November to the end of April,
and in July, August and September. Placed in the order
of their highest monthly averages throughout the year,
{hese common Diatoms are as follows :—

March—Coscinodiscus.
May—Chaetoceras, Thalassiosira and Lauderia.
June—Riizosolenia and Guinardia. |
November—Biddulphia.
On the whole, the regularity of occurrence and of
waxing and waning throughout the year, rather than
differences from year to year, 1s what strikes the observer

as of primary importance.

Biddulphia mobiliensis and

B. sinensis. 7
Both the species of Biddulphia which we have been
obtaining in quantity during recent years (B. mobilensis
and B. sinensis) occurred in October this year, and even
occasionally in September, an unusually early appearance
for B. sinensis. Fig. 1 shows a typical example of a
plankton containing abundance of both species, the
longer and relatively narrower forms in the figure being
B. sinensis, and the shorter, nearly square forms,
B. mobiliensis: there are, of course, other differences
which are not seen clearly in that figure. In the more
enlarged micro-photograph (fig. 2), a points to a typical
B. mobiliensis as seen in our district, and 6 to an

oe nt ici rah

SEA-FISHERIES LABORATORY. 207

Fig. 1.

208 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

example of B. sinensis. Our B. mobiliensis undoubtedly
approaches the form ‘“‘ regza,’’ regarded as a distinct
though allied species by Ostenfeld (Medd. Kom.
Havunders., Plankton, Bd. I. 6, 1908). Gran, in the
Diatomacea of the Nordisches Plankton, unites these two
forms as B. mobiliensvs.

In our district B. senensis is of more elongated form
than is shown in Ostenfeld’s figures. Most of our
specimens of B. sinensts are very distinct and easily
distinguishable from the mobdliensis-regia group by
the shape and the position of the spines, but we have
found one or two specimens during this last year where
one end of the cell bore the characters of sanenszs, while
the other had the appearance of mobiliensis. Until,
however, we get further specimens we do not propose to
base any opinion as to the species upon this possibly
abnormal form. We are watching the fresh material of
B. sinensis carefully in the present year (1912), and may
return to the subject in our next report.

DINOFLAGELLATA.

The monthly averages for Ceratium and Peridiniwm
throughout 1911 were as follows :—

1911. Ceratium | Peridinium 1911. Ceratium | Peridinium
tripos. spp. tripos. spp.
January ... 3,402 44 SY? screens 17,942 864
February ... 1,815 0 August ...... 1,385 125
Marehy 227 1,820 0 September 8,478 33
April ics ccnsce 505 2 October... 8,422 12
May eo. cae: 9,131 6,522 November... 6,382 47
UMC. Sucere 28,811 14,335 December ... 4,866 11

We have taken, in the case of Ceratium, the
familiar form commonly known as C. tripos, without
discriminating between the sub-species or varieties which

SEA-FISHERIES LABORATORY. 209

have been described. The other common species,
C. fusus and C. furca, are also present, although they are
not so abundant as C. tripos.

Under Peridinium, in these reports in the past, we
have united several different forms, and we do so again
in the table above and the one that follows to preserve
continuity; but we shall give a further statement below
as to the sub-genera and species of Perzdinium that occur
in our material.

Looking at the figures from which the table of
monthly averages has been compiled, we find that the
numbers in the case of both Ceratiwm and Peridinium
begin to get larger towards the end of May, and reach
their highest in June and early July. At that time of
year (end of May and June), also, we find that all the
common species are represented in all the nets used, as
the following record of two adjacent hauls demonstrates :

May 29. June Ist.
Species recorded. | Coarse. | Fine. | Vertical.|| Coarse. | Fine. | Vertical.
Ceratium furca...... 300 9,100 200 3,200 | 10,000 2,000
a fusus ...... 100 1,000 50 1,000 2,000 1,500
9 tripos...... 4,500 3,500 150 8,800 | 10,000 2,000
Peridinium spp. ...| 6,600 | 10,000 250 10,000 | 20,000 2,500

Although the above are representative hauls, they
are none of them very large ones. We add the following
records of greater numbers of C. tripos taken in single

hauls :—
POE LOUN.~ f1NO MOUS ii... sc00venscdsenntaewaxe 30,000
, oY oti, Fee ee AD cata Sones dunes eaten es 30,000
: Pee PODS CCOBTSO, MOE! icc tessscssescietsescuss 36,400
; July 15th, pment eis 76,600

Sept. 8th, pat aD coved ike: caeuewus 57,000

910 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.
Irish Sea Species of Peridinium.

Mr. Riddell has drawn up for us the following list,
with diagnoses, of the species of Peridinium we have
found, so far, in our district.

Genus Prripintum.—Testa divided into plates;
precingular plates seven, bottom plates two. :
Sub-genus [.—PRoTOPERIDINIUM: Girdle nearer apex on

right side; usually no antapical horns; spines may

be present. 3 :

(1) P. orbiculare, Paulsen; globular, small; no

spines and no apical horn; girdle almost equa-
torial. L. 0°04—0°045 mm. Rare in Irish Sea.
(See fig. 3, No. 1.)

(2) P. ovatum, Pouchet; depressed; apical horn
not prominent; two antapical spines. L. 0-062
mm., greatest breadth 0084 mm. Not un-
common. (Fig. 3, No. 2.)

(3) P. stent, Jérg.; pyriform; apical horn distinct;

two antapical spines, each winged. L. 0°045—
0°052 mm. Rare. (Fig. 3, No. 3.)

SEA-FISHERIES LABORATORY. 911

(4) P. pellucidum, Bergh; ‘pyriform; girdle at right
angles to long axis; three antapical spines.
L. 0'045—0°068 mm. Rare. (Fig. 3, No. 4.)

Sub-genus I].—Evreripinium: Girdle nearer apex on
left side; antapical horns often present.

(1) P. depressum, Bailey; girdle oblique to long
axis; horns subequal, hollow. L. 0°152—02
mm. Commonest form in Irish Sea. (Fig. 3,
No. 5, A and B.)

(2) P. dwergens, Khrenberg; girdle at right angles
to long axis; inner side of antapical horns with
protuberances. L. 0°08—0°084 mm. Rare.
(Fig. 3, No. 6.)

(3) P. conicum, Gran; girdle at right angles to long
axis; antapical horns short and without pro-
tuberances; left border of longitudinal furrow
straight. L. 0°048—0°06 mm. Rare. (Fig. 3,
No. 7.)

NoctTILuca.

The distribution of Noctiluca throughout the year,
in 1911, was unusual in two respects—(1) in its presence
in fair quantity during many months (ten), and (2) in
having no marked maximum in summer and autumn. It
‘seems to have remained in Port Erin Bay over the winter,
and was caught during the early months of the year in
diminishing numbers—up to 1,200 per haul in January,
300 in February, and 100 in March. It is not recorded
in April and May, but re-appears in small numbers in
June, reaches 1,200 in the best haul in July, a few in
August, and 1,800 in September, 2,000 (the top figure) in
October, fewer in November, and 1,600 in December.

Hauls of between one and two thousand occur at
such diverse times of year as January, July, September,
October and December.

212 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

COPEPODA.

On taking out again the records of occurrence of the
nine commonest species of Copepoda, we get the following
results : — |

Calanus helgolandicus.—As usual, Calanus was
present in small numbers throughout the year, but
became much more abundant in summer, June to
October, and especially in July, when two of the
customary large swarms appeared—one on July 4th and
the other on July 18th. The numbers, however, were
throughout smaller than in 1910.

Pseudocalanus elongatus.—This is one of the most
abundant species, and is present in quantity at Port Erin
all the year round. The numbers begin to get progres-
sively larger in April, May and June, to a maximum in
July, after which they decrease irregularly through
August and September. ‘The greatest haul (54,350 on
July 3lst) is more than twice as large as the top haul
of 1910. |

Oithona similis.—The commonest Copepod in Port
Erin Bay throughout the year is again Ozthona similis.
The greatest haul is 225,450 on July 18th, a very large
number for a Copepod, in our standard hauls, and nearly
twice as much as the largest (126,700 at Station I on
August 20th) of the previous year. In general, the
record is the same as before.

Temora longicornis.—This is a summer form, and
this year it was only abundant in June and July. It has
a remarkably symmetrical distribution, forming a simple
normal curve, rising from an average of 1 per haul in
January, having the maximum in summer, and sinking
to 2 per haul in December. The average for June is
4,675 per haul and for July 4,706, and the two largest

— es

SEA-FISHERIES LABORATORY. 213

hauls are 20,000 on June 3rd, and 17,900 on July 4th.
Temora, with the same distribution throughout the year,
was more abundant in 1910 and had its maximum at the
end of July.

Paracalanus parvus.—ULast year’s remarks in regard
to this species apply again. The maximum is again in
September, and the largest hauls are 61,930 on August
24th, and 32,390 on September 29th.

Acartia clausi1.—This species again only reaches high
numbers in summer, and this year has its maximum in
August—the only really large hauls were 59,360 on
August 24th and 42,380 on October 19th.

Anomalocera patersoni.—This oceanic form was
exceptionally rare at Port Erin in 1911, and was absent
during most months of the year. A few stray individuals
were present in June and July.

Centropages hamatus.—This is a summer form
appearing in April, becoming more abundant in May,
with a low maximum in June, and then dwindling
gradually to September.

Microcalanus pusillus.—Uast year’s record again
holds good for this species. The maximum in 1911 is in
December, and the minimum in July, August and
September.

Amongst other rarer Copepoda, Huterpina acutifrons
is a winter form which occurred in fair quantity in
November, December and January, and in no other
month. Jszas clavipes occurred in May, June, August,
September and October, the two largest hauls being 130
on May 18th, and 156 on September 13th.

Omitting these latter, rarer, species, the monthly
average hauls in Port Erin Bay for the above nine, more
important, species of Copepoda are as follows : —

914 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

3 ; : -

a g | 8 : :

a on oO lm as}

1911 = 5 B SF aS) od & a

s & s ° S s = 3S S

=| S) a q ~ cS) 3) fo)

2 5 q = S a a s 5

5 Ay HB O < < fo) a =
Jan. 4 370 1 ) 0 60 2,589 99 14
Feb. 8 460 0 0 0 1 1,993 61 38
Mar. ... 1 476° 63 | 0 0 17 428 23 5
April: <:: 4 1,554 305 a 0) 197 675 7 ey
May .. 15 1,501 282 | 42 0 1,627 5,647 39 ll
June... 85 4,168 4,675 | 81 i 4,858 | 11,745 326 32
July ...| 339 | 12,012 4,706 | 45 1 5,376 | 52,393 667 0
ANG Te 32 8,916 435 | 34 0 14,049 | 20,755 | 13,372 0
Sept. ... 68 4,129 69 | 28 0 4,234 9,000 | 14,100 0
Och. 32 60 2,332 Pail i 0 6,181 5,024 5,170 12
Novi =: 3 3,056 3 0 0 222 | LiZonZ 5,506 on
Dees. 5: 4 1,184 2 0 0 51 6,587 1,860 89

It must be remembered that the above figures are
only the monthly averages, per haul, and that many of
the individual hauls in each month were much larger, as
we shall show below. Anomalocera, as it appears in the
above table, would not be worthy of record were it not
that it has assumed more importance in other years, and
may do so again, and consequently it is well to preserve
this year’s record for comparison. .

If we arrange the six most abundant of these species
of Copepoda in the order of their abundance with their
total numbers—and add the totals for the two previous
years—they are as follows :—

1911. 1910. 1909.
OuUhona svmilis Vs. .orsceee aici ceeee 1,155,108 872,678 465,066
Pseudocalanus elongatus ......... 365,983 368,326 309,973
Paracalanus.Darvus ....2.¢-60--. 4 351,088 217,633 54,120
ACOLUG CLAUS Us. ..2 o oemeeeeeeeeeeene 323,633 340,631 63,373
Temora longicornis “ne seen see. ee 106,359 147,043 62,659
Calanus helgolandicus .........++. 5,843 15,481 21,412

This list shows that the only difference in the order
is that this year Paracalanus parvus comes above Acartia

claust, while it was below in 1910 and 1909. The

le he ad

SEA-FISHERIES LABORATORY. 215

progressive increase in the numbers of Ozthona similis
during the three years is noteworthy.

The greatest single hauls of Copepoda (taken with the
ordinary ‘‘ coarse’’ net) in Port Hrin Bay, during 1911,
were as follows :—

Species. Number. Date.

DU OUE SURMISE... 020... 0cinc eee cenans 214,000 July 18
73,200 eee |

68,000 Aug. 3

51,300 5 ok

Pseudocalanus elongatus ............ 46,500 July 31
34,350 Aug. 24

PANGCIANUS PATVUS ....... 5200000000 61,300 said dee
32,300 Sept. 29

TAOTTTRO CIUUSY oc sceseeccseccsvceseesuss 59,000 Aug. 24
42,100 Oct. 19

MCMEOR VONGUCOTNIS 2.2 20..s0eeceneess 17,000 June 3
, 17,000 July 4

Calanus helgolandicus .............4: 1,800 was

Some of these are very large hauls indeed for
Copepoda.

If we arrange all the species of Copepoda (15)
recorded on our forms in 1911 in two series, according
as they are supposed to be ‘‘ Oceanic ’”’ or “‘ Neritic,’’ we
get the following result : —

OcEANIC. NERITIC.
Calanus helgolandicus. Temora longicornis.
Pseudocalanus elongatus. [Centropages hamatus |
Oithona similis. Euterpina acuttfrons.
Paracalanus parvus. Isias clavipes.

[Acartia claust| Parapontella brevicornis.
Anomalocera patersoni. Labidocera wollastoni.

Microcalanus pusillus.
Metridia lucens.
Candacia armata.

A few of these are undoubtedly oceanic and neritic
respectively, for example Calanus helgolandicus is
typically oceanic, while /sias clavipes is a shore form;
but other cases are more doubtful. One would expect an
oceanic form which only invades a shore area on
occasions to have a more or less periodic distribution, but

916 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Pseudocalanus elongatus and Oithona similis, usually
regarded as oceanic, are present in quantity in Port Erin
Bay all the year round. Again Temora longicornis, a
reputed neritic species, is at Port Erin a summer form
only abundant in June and July—a distribution which
might be supposed to be characteristic of the immigration
of an oceanic species. It must be remembered, however,
that periodicity, and absence or great reduction in
numbers during part of the year may be due to the normal
life-history of a neritic species in situ and need not
indicate invasion from outside.

The fresh evidence this year tends merely to confirm
us in the opinion expressed in last year’s report*, to the
effect that—‘‘ It is improbable that all planktonic species
are either oceanic or neritic. It may well be that some
species are intermediate in character and habitat, over-
lapping and intermingling with both, and liable to be
placed sometimes in the one category and sometimes in
the other. Then again, there may be some species which
are cosmopolitan, or ‘ Panthalassic,’ as we should prefer
to call it, occurring both in the open oceans and also in
the shallower coastal waters of some parts of the world.”’

CLADOCERA.

This summer group ranged in 1911 from March 2nd
to September 29th. Podon (with 3,500), and Hvadne
~ (with 6,460), both reached their highest point early In
September; but Podon also showed large hauls in June
(1,320) and August (1,350), and Evadne in June (6,000),
July (2,400), and August (2,550). These numbers, it will
be noticed, are distinctly larger than those for 1910.

* Lancashire Sea Fisheries Laboratory Report for 1910, p. 107.

‘

SEA-FISHERIES LABORATORY. 217

SAGITTA.

This ‘‘ Panthalassic’’ species (S. bipunctata) usually
considered to be an oceanic form, was again present in
Port Erin Bay in every month, with the maximum in
summer (June), and a secondary increase in late autumn
(November). It became more abundant in November after
stormy weather had set in.

The monthly average hauls are as follows : —

Jan. Feb. Mar. Apr. May June July Aug. Sept. Oct. Nov. Dec.
9 a 4 7 66 244 64 30 15 28 59 9

The largest hauls are 206 on May 29th, 620 on
June Ist, 1,300 on June 3rd, 250 on June 15th, and 412
on November 24th.

A haul of the shear-net five miles off land, at about
10 fathoms, on April 14th, gave 301 large specimens of
Sagitta, although the surface nets worked at the same
time caught none, and the medium Nansen hauled
vertically from 20 fathoms showed only 4. This result
suggests that at that time although not present at the
surface Sagitta was abundant a few fathoms below.

OIKOPLEURA.

This form (QO. dioica) has again the same sort of
distribution curve throughout the year. It is present in
every month, but the monthly average hauls only run to
thousands from May to October, inclusive. The largest
hauls are over 34,000 on May Ist, and over 15,000 on
October 19th.

Various LARVAE.

Cirripede nauphi and other larval forms do not show
anything very striking in their distribution this year,
and so do not call for individual treatment. It will

918 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

suffice to put on record a few of the more exceptional
hauls. Nearly 130,000 young Polychaet larvae were
taken in Port Erin Bay on February 17th, and over
09,000 on March 11th, 8,000 Lamellibranch fry on May
4th, 11,000 Gastropod larvae on July 18th, over 2,000
Kchinoderm larvae on August 7th, over 5,000 on
September 4th, and 4,700 on September 11th, over 4,000
young Medusae on September 4th, and over 5,000
‘“ Mitraria’’ on December 26th. In the later part of
the year the following large hauls of Lamellibranch
larvae were taken: 35,000 on September 25th, 17,000 on
October 2nd, 19,000 on October 27th, 15,000 on November
1st, 11,000 on November 10th, and 22,000 on December
26th. It is probable that the large scallop bed (Pecten
opercularis) lying outside Port Erin Bay to the North
accounts for many of the young Lamellibranch fry that
are so abundant from time to time.

Fisu EGGs.

Rockling eggs continue to be abundant, and have an
extraordinarily wide range through the year, as they are
only absent in October and November. The maximum is
in March, with an average of 49 per haul. The eggs of
one of the species of Rockling occurred in a sample of
Plankton collected on December 29th, which is unusually
early, as these eggs are not often found in our seas before
the end of January. The ‘‘ mackerel midge’’ (a little
narrow silvery fish about an inch long), which is a young
stage of one of the Rocklings, is of economic importance
as it sometimes forms a considerable part of the food of
the Mackerel visiting the Irish Sea in summer. The
young Rockling live near the surface and are frequently
taken in the tow-net between June and August.

SEA-FISHERIES LABORATORY. 219

The other fish eggs range from January to August
inclusive, with a maximum of about 20 per haul in the
bay during March and April. Out at sea during April
the average per haul is about 36, the largest hauls being
62 per net on April 10th and 63 per net on April 27th.

A haul of the large Nansen net on April 12th secured
1,059 fish eggs, and 2,470 on April 24th. Both these
hauls were outside the bay, and the eggs caught were as
follows :—

. April 12 April 24
00 5 ee 750 650
Ee cis a tjejo cad one ssiewanea's 25 300
ae ee nines Socio eiscacisiews 230 1,500
PMD ein sda on creSecvigencie ese acs owe 50
“du LLG ee a 0 20
RIN Meee cece crsee cestesrncens ees 4 0

COMPARISON OF NETS AND CATCHES.

There is little, if anything, to add as to the catching
power of the different nets to what we have written in the
previous parts of this report. It may just be worth
noting that :—

In our experimental hauls from the yacht the coarse
net consistently caught from twice to ten times as much
as the fine, both in spring and autumn.

The two nets fitted with Otter boards and towed well
forward in the ship, so as not to be affected by the
propeller, agreed fairly well in their results with one
another, and with the fine net at the stern. There is
nothing to add under this head to what we have already
said.

With the small (35 cm. diameter) closing Nansen net,
we find that when two hauls are taken together, one
from 20 to 10 fathoms, and the other from 20 fathoms to
the surface, the latter in many cases gives, as would be
expected, the larger volume of plankton, such as, April

220 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

18th, Station ITI, 20-10 = 0°9:c.c., 20-0 = 1°5 ¢.c.5 but in
other cases again we find that the two hauls are roughly
equal in volume, indicating that the bulk of the plankton
is in the zone below 10 fathoms from the surface, as for
example, April 11th, Stat. I., 20-10 and 20-0 fathoms,
both = 0°6 c.c. The medium Nansen (50 cm. diameter)
hauled on the same occasion from 20-0 fathoms gave
16 c.c.; and from this to three times seems the usual
proportion between these two nets when hauled vertically
together.

The medium Nansen, and also the large Nansen
(100 cm. diam.) were sometimes towed horizontally, like
the shear-net, when it was desired to obtain not a standard
haul for statistical comparison, but a large bulk of
material for some special purpose.

In comparing the hauls, taken with exactly similar
nets, in the bay on the one hand and in the open sea
from the yacht on the other, we find that both in spring
and in autumn the hauls from the sea outside were on
the whole larger than those from the bay. The difference
is usually not very large, but the records seem to indicate
that the amount of plankton (Diatoms, Dinoflagellates
and Copepoda) in the open sea is rather greater than that
in the bay during April, August and September.

WEATHER CONDITIONS.

The weather during 1911 was certainly exceptional,
and this might be expected possibly to have some effect
upon the amount of the plankton. The spring was cold,
and the latter part of the summer (especially August) was
unusually dry and warm. Professor Bassett, in his
article on the Hydrography of the Irish Sea in 1911,
associates these weather conditions with an unusually

SEA-FISHERIES LABORATORY. 221

early time of arrival and greater strength of the Gulf
Stream Drift in our area. After two successive years of
a late and weak Gulf Stream Drift associated with wet
and gloomy summers, we had in 1911 an early and
unusually strong invasion of Atlantic water in the Irish
Sea, followed by the exceptionally dry and sunny
summer. Under these unusual conditions we have
decided that this cannot be regarded as a normal year,
and that it is desirable to carry on the observations a
little longer in the hope of obtaining a greater uniformity
of results. |

We insert here for reference the chart (fig. 4) of air
and sea temperatures for 1911; and, for comparison, the
similar chart (fig. 5) of the previous year—both made
from the Port Erin Biological Station records by Mr.
H. C. Chadwick. It will be noticed that, compared with
1910, the sea-temperature in 1911 was lower in spring,
below 43° F. in the first week of April, but reached a
higher point in summer, over 59° F. late in August.
There was an almost uninterrupted rise in temperature
for five months, from March 25th (the lowest sea-tem-
perature of the year, 42° F.) to August 26th. In the air-
temperatures the highest weekly average was 63° F., and
an unusual number of records are above 60° F., one week
in June, two in July, and four in August. In 1910 only
one week (August) was over 60°, and in 1909 the highest
weekly average was 58° F.

SUNSHINE.

We are only concerned with the hours of sunshine in
so far as the record seems to show any co-relation with
the plankton. The summer of 1911 was a notable one
as regards amount of sunshine. In May, June, August,
and September, the hours of sunshine at Port Erin were

922 TRANSACTIONS LIVERPOOL BIOLOGICAL, SOCIETY.

VAN. FEB. MAR. APR. MAY. JUNE JULY. AUC. SEPT OCT NOV. DEC.
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SEA-FISHERIES LABORATORY. 923

considerably above those of the corresponding months of
1910. Some other months were a little over, July was a
little under, but August had no less than 194 hours
recorded, as against 80 hours in 1910, and about 107 as
the average of the previous four years. It is clear,
however, that this summer and autumn sunshine can have
no effect upon the spring phytoplankton. It is, as we
showed last year, the March sunshine that may be co-
related with that; and any effect of the enormous increase
in the August sunshine must be looked for in the autumn
and winter plankton—or possibly even in that of the
following season. We have seen above that some
elements of the plankton were unusually large last
autumn, such as the Diatoms. This is well shown by
the Diatom monthly averages per haul in the two years:

1910. 1911.
EIR ss asin a 0 vor v0 850 1,998
September ............... 676,823 928,501
oor 553,601 4,742,791
Movember ..2............ 100,262 506,729

In both years the autumnal increase begins in
September, and reaches its maximum either in that
month or in October and falls off in November; but in
1911 the maximum is about nine times as great as in the
previous year.

Some of the groups of the zooplankton, notably
Cladocera, Polychaet larvae, and Lamellibranch larvae,
also show larger numbers this year in autumn and early
winter than has been usual. Whether this is to be
regarded as a result of the great increase in the amount
of sunshine last summer, and if so whether the connection
is direct or is in the case of the animals a result of the
increased number of Diatoms, we are not prepared to say.

224 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Amongst the Diatoms, in spring, species of
Chaetoceras were in much greater abundance than usual
this year, and in autumn Biddulphia, Thalassiosira and
Chaetoceras were greater in amount than in any of the
previous years of our study at corresponding times.

In CONCLUSION.

Fach additional year’s work tends to confirm us in
our view that, although there is a natural sequence in the
distribution of the plankton throughout the year, and
although there is a certain constancy in the maxima and
minima for particular groups and even species, still the
sequence is liable to disturbance and the maxima are
affected both in time and in amount by surrounding
conditions. This leads to the numerous variations which
we have had to record from year to year.

Moreover, even when some constituent of the
plankton is most abundant, its distribution may be
irregular, in streaks or in patches, so as to destroy any
such uniformity as would justify small samples being
taken as representative of wide areas.

We are not yet prepared to make the promised
collation and comparison of our records for the five years,
that was referred to in the last Report. We shall hope
to do so in a final part next year. —

J
.
:
(
i
|

SEA-FISHERIES LABORATORY. 225

THE PLANKTON ON THE WEST COAST OF
eoummanl) IN’) RELATION TO THAT OF
THE IRISH SEA.—PART II.

By W. A. Herpman, F.R.S., and Wm. Rippetz, M.A.

InrTRopuctory Note.

In last year’s Report we pointed out that our know-
ledge of the plankton of the western coasts of the British
Islands is incomplete, by reason of a great gap extending
from the North of Scotland down to the Irish Sea—a gap
in our knowledge which neither the International observa-
tions on the one hand nor those of the Irish or Scottish
Authorities on the other seem to fill up. With the view
of obtaining data, which might in part at least bridge this
gap and possibly throw lght upon the question of the
seasonal changes in the plankton of the Irish Sea, one of
us has for several years, during the summer vacation, »
taken plankton hauls, both vertical and horizontal, from
his yacht at various localities amongst the islands and sea
lochs of the west of Scotland as far north as Portree in
Skye, and as far out to the west as the Island of Barra.
In our paper last year we discussed these data, and were
able to show that the state of affairs in these Scottish seas,
at that time of year, is somewhat different from that in
the Irish Sea. At some spots in the Hebridean seas, for
example, large phytoplankton hauls may be’ taken in
July, at the time when in Manx waters the hauls are
comparatively small, and are composed of zooplankton.
It thus becomes of fundamental importance in connection
with local sea-fisheries problems to determine more accu-
rately the relation of the Irish Sea plankton to that of
neighbouring waters to the north, to the south, and in the
Atlantic outside. Such information may enable us to

226 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

estimate, for example, how far our local seasonal changes
in the plankton are due to migrations or invasions from
outside waters. |

As the region to the south of the Irish Sea, including
the English Channel, is being thoroughly investigated
under the scheme of the International Council, and the
necessary data will therefore naturally be available from
that source, it becomes all the more important te do all
that is possible to get information in regard to the con-
dition of the plankton in the seas off the north of Ireland,
and the west and north-west coasts of Scotland.

Cruise or 1911.

During this last summer, July and August, 1911, we
were able to devote a longer time than in previous years
to a more detailed survey, from the yacht “ Runa,” with
both bottom and surface nets, of a considerable area of the
west and north of Scotland. Our observations extend
from the Irish Sea as far north as Noup of Noss in Shet-
land (from about 54° N. lat. to 60° N. lat.), and as far
west as Castle Bay in Barra. They include 152 observa-
tions of the sea-temperature, and 142 of the salinity, and
not only sample the water at a number of points lying off
the Coast of Scotland, but also give us a _ series
of observations across the northern entrance to
the Irish Sea, as follows:—On August 22nd,
when crossing from the south end of Cantyre
to the north of Ireland, a series of nine temperature and
salinity observations were taken, one every hour during
the most important part of the traverse; and on the
following day when crossing from Larne in Ireland to
Port Erin, another series of eleven hourly observations
was taken.

During these two months (July 7th to August

SEA-FISHERIES LABORATORY. vad

23rd) the temperatures varied from 112° C. to 178° C.,
and the salinities from 1°018 to 1°:0276, the latter
reading being a high salinity for British seas. It was
recorded on August 12th in the open sea to the east of
the Shetlands, but nearly as high a reading was obtained
off Fair Island, off North Ronaldsay, and elsewhere in the
Orkney seas, and 1°027 was obtained on July 13th and
14th off Canna and Rum on the west of Scotland.

PHYSICAL OBSERVATIONS.

The full list of our observations on temperature and
salinities 1s appended to this paper. The list gives
the dates, localities and fimes of the observations,
the temperatures Centigrade, as taken in a cooled, clean
canvas ship's bucket, rinsed and dipped from the forward
part of the ship, and tested with the thermometer without
delay. The next column gives the observations as taken
at the time with a Kiel ardometer immersed in a cylinder
of the same water from the bucket, the cylinder having
been rinsed with a sample of the water before being filled.
The sixth column gives the specific gravities at 17°5° C.,
as a result of reducing these readings at the given
temperatures by means of Knudsen’s Hydrographical
Tables.

On looking at the first column in the list it is clear
that some of the temperatures are merely due to temporary
conditions of the locality—such as 17°8° C. at Loch
Scavaig, in Skye, at 10.30 p.m., after a hot calm day,
when the high temperature was no doubt due to the effect
of the sun on the neighbouring rocks during the afternoon.
On the other hand, several of the series of temperatures
clearly indicate tracts of colder water. For example, in
the North Channel and round the Mull of Cantyre, our
readings lie between 11° and 12°, which is lower than the

228 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,

temperature of the water both to the south and to the
north. Then again, the following day we had water below
12° all the way up the Sound of Jura, while at Oban, and
in the Sound of Lorn and Sound of Mull the temperatures
were mostly between 13° and 15°. Off the north-west
mainland to Cape Wrath the temperatures are between

12° and 13°, while further no1th in the Orkneys, cff Fair

Island, and off Sumburgh Head in Shetland, 14° and over
isreached. Higher temperatures, about 15°, were recorded
south of Skye around the Small Isles later in August,
and again in the Irish Sea when returning to the Isle of
Man. It is curious that the highest temperature (16°1°)
recorded that day (August 23rd) in crossing from the
North of Ireland to Port Erin was found in mid-channel.
This observation was repeated to ensure that no mistake
had been made.

The column giving the araometer readings shows
that compared with water at about ...26 in the Irish Sea
the North Channel has 26°5, while the neighbourhood of
Oban, the Lynn of Morven and the Sound of Mull show
lower readings of from 23 to 25. Round the outside of
Mull to Staffa and Iona the water is fairly constant at
from 26 to 26-4. In places round the Islands south of
Skye, and again in the Outer Hebrides round Barra, 27 is
reached. The water off the north-west coast of Scotland
is mainly between 26 and 268; while further north in
the Orkneys, off Fair Island, and in the Shetlands 27 and
over is reached. The highest reading is 27°6 in Bressay
Sound on August 12th. Although our readings are, we
believe, comparable with one another for the purpose of

such contrasts as we have made above, we wish to state

that we are inclined to regard them all as uniformly too
low, and requiring a correction applied (which we have not
yet determined) before they can be used for comparison
with other series of observations.

SEA-FISHERIES LABORATORY. 229

PLANKTON OBSERVATIONS.

The plankton hauls, taken as frequently as possible
simultaneously with the temperature and salinity obser-
vations, were in all cases made with the same nets and by
the same method so that the various gatherings might be
as nearly comparable as possible. All the vertical hauls
were made with the smaller Nansen closing net, of No.
20 silk, and having a mouth 35 cm. in diameter. Surface
hauls were sometimes taken at the same time, with
ordinary open surface tow-nets made of the same silk
(No. 20) as the Nansen net and of approximately the
same size of mouth. The Lucas sounding machine, fitted
with 200 fathoms of pianoforte wire, was used in all cases
in taking the vertical hauls—down to a depth of 135
fathoms.

_ The list given below at page 230 shows the complete
series of these plankton gatherings, which will be found
to represent most of the localities dealt with in our paper
in last year’s Report, and we have, moreover, extended
this year the observations considerably further to the
north—from the Island of Skye to the Shetland Isles.
We do not propose in this year’s Report to make a
detailed analysis of these hauls, as we hope to be able to
repeat a number of the observations in the coming
season; but in comparing even the brief descriptions
given in our list with the characteristics of the plankton
at the same localities in last year’s Report, we notice
certain differences which we desire to point out. For that
purpose we shall group this year’s 70 gatherings into
seven areas comparable as far as possible with those of
last year’s Report, and shall indicate briefly the nature
of the plankton in each and note any differences that are
seen.

TRANSACTIONS LIVERPOOL BICLOGICAL SOCIETY.

230

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231

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LABORATORY.

SEA-FISHERIES

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232 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

COMPARISON WITH PREVIOUS YEARS.

We have no data this year representing the Clyde
Sea-Area, but can deal with all our previous localities
north of Cantyre, as follows :—

Firth of Lorn.

Half a dozen hauls, ranging between Sheep Island
and the Lynn of Morven—mixed hauls, but on the whole
more zooplanktonic than in previous years. The Diatoms
are mainly neritic. A haul taken off Lismore on July
10th is the only purely phytoplanktonic gathering in this
series, and contains about fifteen millions of Chaetoceras.
The rest of the hauls from this district were taken a
couple of weeks later between the 24th and 26th of July,
and it is quite likely that their more zooplanktonic
character is due to the seasonal disappearance of the
phytoplankton. 7

North End of Sound of Mull.

Hauls at Tobermory show much the same characters
as in previous years, but one off Ardmore, on July 11th,
presents a striking difference as it is almost a pure zoo-
plankton, the Diatoms which were present to the number
of over 40 millions per haul the previous year being
represented now by only a very few Chaetoceras and
Coscinodiscus, about 1600 in all. The details of the 1910
and 1911 hauls at this locality are given below for com-
parison. It was here, this year, that we first met with
the Pteropods (Limacina retroversa) which were so
abundant from this point onwards up to the Orkneys.

Hebridean Sea and Round the Small Islands.

Fifteen hauls represent this area, and on the whole
they are more zooplanktonic than in previous years, and
show a change in the nature of the few Diatoms present,

SEA-FISHERIES LABORATORY. 233

most of which are of an oceanic type (such as, Chaetoceras
densum and Rhizosolenia alata). The Pteropod Lima-
cina is present in nearly all cases in abundance (up to

about 30,000 in a haul).
Sound of Sleat.

Five hauls in Loch Hourn and Loch Nevis showed
very much the same characteristics as in previous years.
North of Narrows of Skye—lInner Sound and Lochs.

Nine hauls were taken between the northern part of
Skye and the mainland, and fall into two series, those in
the lochs being phytoplanktonic and mainly neritic,

while those in the opener water of the inner sound were

mainly a zooplankton. ‘So far as they are comparable
these give much the same evidence as in previous years.

This brings us to the furthest point north reached in
previous years, but we add the characteristics of the
remaining localities for comparison with those further
south.

North-West Coast from Ru Rea to Cape Wrath.

Seven hauls taken on August 15th and 16th show
mixed gatherings, with the exception of a haul in Loch
Inchard which is a typical phytoplankton containing
nearly 50 millions of Diatoms (the details of this haul are
given below). Some are more zooplanktonic than others,
but in general there are far more Diatoms here than in the
Hebridean seas south of Skye.

From Cape Wrath to the Shetland Isles.

Four hauls, two phytoplanktonic and two mixed
gatherings containing both animals and plants, of both
neritic and oceanic species, show at least that the Diatoms
had not disappeared by the middle of August at this
furthest north point in the British Seas.

234. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

NoTEWORTHY SPECIES AND HAULS.

There are a few of the species occurring in this year’s
hauls that are of special interest. In the Diatoms,
Skeletonema costatum, which had been found in great
abundance by George Murray in 1896, in Loch Etive,
and which we only met with rarely in previous years,
occurs more plentifully in our gatherings this year in the
Mull district. Amongst new records we have Netzschia
clostertum (Loch Scavaig, Skye), Chaetoceras atlanticum
(Moussa Sound) and Pyrocystis lunula (Loch Hourn and
Loch Nevis). |

In Upper Loch Torridon, on July 19th, with water
of very low salinity (1°018)—due no doubt to mixture
with fresh water from the land—we got two remarkable
hauls (coarse and fine nets) containing, along with a

Upper Loch Torridon, 19/7/11. Fine Net. Coarse Net.
Calanus ’ helgolandicus .............s05.00.000. 6 —
Pseudocalanus elongatus§ .................. x —
(Nest © Wonie..ceSea Be seen ti eons cose ener neeeeEne 10,000 15,700
TOs) nip eros peer bene aondser cocsece cedsocdccoces — 9
Oikoplemra.?/25-0%.. CORR Geto aeaceenee 136,500 42,000
Mam Git dase. S950. sic saan eee one see emeeence 420,000 388,500
Ceratimimir furca, Ju. ..iccses smote eaten dome cnaens x _
- CETPIOSS) -..hiceccesisern sor casemate gen — 5,250
Pericl iminm SOPs... .2- sec. ssocehemencecetiseten as ~~ —~2,052,500 351,700
DinOphiysis) SPs4q Gero se-wssdseeeor saeco nee 189,000 21,000
Chaetoceras boreale yy .caereete eee ce ose eters 1,470,000 1,454,000
183 COUSELICHUM™ oo .ce ese ctincdeco es 1,102,500 141,700
os debiles g.c28 sateen easeceess — 241,500
~ AECIPIONS WOCe whos eae acess e eee 2,782,500 5,076,700
os S10) Oo Ga OO ACE Se escn cana he mame 16,117,500 2,709,000
Nitzschia delicatissimay assesses sete: ee eee. 334,365,000 328,125 000
ae seriata, ¢aceetesctesccsans cata sleses 73,500 36,700
Rhizosolenia semispina ................05+++ 346,500 871,500
Skeletonema costatum ...............-..+++ 84,000 84,000
Thalassiosira decipiens .................066- 378,000 —
SEAVIGS: oe cwesoncpeestortvsesies 462,000 771,700
number of the usual marine Diatoms, an enormous

quantity of Wtzschia delicatissima which. we estimate at
well over three hundred million individuals in each net.
In this case the size of the mesh seemed to make so little
difference in the nature and amount of the catch that we

SEA-FISHERIES LABORATORY. 935

think it well to give the details of these two hauls here as
a record of this unusual occurrence.

One of the most noticeable features of this year’s
gatherings was the abundance and wide distribution of
the Pteropod Limacina retroversa. After we first came
upon it at Tobermory in July, it was practically
universal in its presence and extended from the north of
Mull up to the Orkneys and Shetlands. It was only
absent in a few of the more sheltered and land-locked
localities. The numbers in a haul vary from about 400
(at Tobermory) to 30,000 (South of Skye and also at Loch
Scresort in Rum).

This Pteropod is a boreal Atlantic form which is
found commonly off the south and west of Ireland—
where it isa common food of the mackerel. Dr. T. Scott
has also found it as a common food for herrings from the
west coast of Scotland. It does not extend up to Arctic
seas; it is rare in the English Channel, does not reach
the North Sea and apparently does not pass up through
St. George’s Channel.

We have selected a few other notable hauls to give
here in detail :—

1. Off Ardmore, for comparison with the previous
year.

Between Canna and Rum, again for comparison.

Loch Inchard, to show an undoubted phyto-
plankton far north in the middle of August.

4. Moussa Sound, Shetland—one of our most
northerly hauls, and again a phytoplankton.

236 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

1. Off Ardmore, N. of Mull.

July 18th, 1910.July 11th, 1911.
86 fathoms.

105 fathoms.

Chaetoceras curvisetum .........:ccssssecees 18,571,450 —
*5 debile ti aanc pe. eatreheead 6,571,450 —
me CEGCIPIONS” <5 Yona csesaecesseeeeese 2,071,440 —
a SPs hile eocsen et duets Be aecacas 20,357,150 1,260

Coscinodiscus radiatus — .............sceeeeee 2,500 420

Dityliimbrightwellii))62.200d/.00¢s.sse0ces ss 625 —

Cutmardia Maccidanea mn e.sdecreenece cs sense 71,425 —

audertasborealis, “etic. ences. w<s0s en. soesoane 357,145 —

Nitzschia seria tan 2 Bysaceccedecst oc « anaoosnees 1,642,850 —

Pleurosigma, SPs». cee s.scrisscn cece esos asec 1,250 _

Rhizosolenia semispina .................000 464,285 —

os SEUISCN A sare dace ens ance ee 17,860 —
a Shra soli nt mas cece eee 190,475 © —
ye stolterfotii «joc. cessescc.ceeees 2,142,860 —

Skeletonema costatum ............-esceees 857,145 —

Tholassiosira: gravida!) sts 0.60.5tds0s dee tes ee 1,428,575 —

‘ NGLAENSKIONMT 964 e teers nes 571,430 —

Peridinriia SO Prcssvaccucwss sees o: es oeceer tence 10,000 1,260

Ceratinm .filrea 2..suiuges sends setae tat deeuae 625 —
53 ETUPOS: She's sine ose ngcnises ace ncoeaeewen 625 840

Pinatas PEE Ska sreess Aneeek se dae ee 13,440 840

IM@GSRG: aoe neces otene caret cet ee as 13 6

WAEUba” tence scoetuce Moree oeeeea tere en mares 25 30

MOMOPtETIS use sete tome esses cores nance — 1

Polychaet larvae (st Acerecce oernastececee «Hs 625 —

Ph ute? ue ite cecs cheb setae enone naan eee 3,600 —_

Gastropod Jarvae asksio0 stnee ce eee 7,500 —

Lamellibranch larvae) occ.ccssecescseeaeseeene 5,000 —

Limaema, retroveraa $225.00. .k Saco ete — 9,360

SEMIZO MOA soican scutes tan cower cee See 6 x

Evadne: nordmanniG’.c.ce-easce-ce sens speeeeee — 525

Calanus helgolandicus .......2......-0.0<---.. 7 > 95 430

Pseudocalanus elongatus 9 ................4. 5,100 2,100

Temora, longicornis ©..sis<<-22ss0.2s55-5cerse os 935 3,150

Oithona similisascs: vessseaees.cea awe eaten ene 625 1,150

Copepoda quves sctnosccecmeo-ctereesceetee chase 1,875 —

INavinplit sy aik nets ea dade cc JES sets Ateeee es 25,625 8,800

Oikopleura, (nse hhienccsaectie ls danecsoeeseeee 3,230 420

This year’s haul (right hand column) shows a some-
what scanty zooplankton, last year’s a very abundant
These two hauls show clearly that the
main difference is not the presence of additional animals
in the zooplankton, but merely the absence of the

phytoplankton.

Diatoms.

== SS

all

SEA-FISHERIES LABORATORY. 237

July 14th, 1910.July 13th, 1911
2. Between Canna and Rum. 128 fathoms. 130 fathoms.

Chaetoceras constrictum ...............0cee0. 5,714,300 —
_ MP MIRCHUIE — <ci0s0tecsiccccnsss 41,714,200 _

+ “SILL. ee ere 5,285,700 —

- MERON. winidviesleeescisitavsadeess 3,857,140 1,050

= Lib. aongesngetooaeapepaanecenosnse 15,214,275 —
WeseHOURCONTACGIATUS .......2.0cesccececes 4,690 420
ATTA ACCIOR,........0c0ccecccnscavacsveces 1,040 —
ICECTELCANIS, 22... .c0cc.ccccccesveccesecces 23,960 —
ITPRGONAPBCEIAUA ...0..200eccccscecccececdecsss 357,140 —
OMIM) Sesh ccinsecceccencscceccsecsecd x —
Rhizosolenia semispina .................000 142,750 840
ss shrubsolii ........ Seats 29,175 —

te solperfothil) 2. .....css2.s0000 17,715 —

f, SEM MIOUMIS .ic.c cesses ese ee a 420

EE MIMEETTIAMISUIS oss 55cs000ssa0cscaaveevecess’vece 1,040 —
se PMECTIBCOIUM, i c065cecceeeccceeess — 200

ss BOVE GIS icBie ss scknstabtweantenevs sien 5,210 3,350
SPIMETTE BID. 02500 0nvccscerccescscccsccsssess 18,750 —
Eo LS 22,920 630
Moire sin as cine csi satancesnceccetsaess 50 _
MRE ate asin ivecesccsaccrcceencaacaaese 6 7
EC MAER MAT VAD. 2. c0ccccrsccseccncsecnsssesens % 200
Rey. putercstessekessareceeccetas 520 —
PARUOPOG AAT VAG’ 5.22. ..0cccccccecssscvscccnees 56,270 —
MAINACIAA TOULOVETSA «= .....0ccesccccscvecncess — 8,400
Calanus helgolandicus .................sesse0 500 520
Pseudocalanus elongatus ..............000. 3,125 2,100
SMUT Secs cccs acs ccdecvcescarecscess 1,215 —-
MN RIITUPRTMEIIIIE, Son cacsccccccceccavcesscssccsesss 4,430 3,150
geo re depicciavssecsiacswececsecsensesees 23,445 9,650
RPM PIOTOTIANINT 2... .cccsnccscccensscscesene oe 200
eh s Jo gic cccvexe tbs snrcdadsanexees 2,080 2,300

This again shows a zooplankton in 1911 at a locality
where at the same time of year in 1910 there was a
marked phytoplankton. The difference 1s due again
simply to the absence of the millions of Diatoms that
characterised the locality in 1910—and the presence of
Pteropods in 1911,

238 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

3. L. Inchard, near Cape Wrath, 15/8/41. Surface.

Chaetoceras boreale: 2. .tevo.cosceecors cen eee eee 378,000

e DEOVO OR Ca ccet oss sneoe eae eee ee 315,000

A COMSPLICHUMA ns aan s-eseceee t eee 11,250,000

e criophilama cco. tech sacoeee aes 2,333,400

Br AG (6) Ov (hai mere ae eee otter at 22,475,000

A GECIPIENS' |. fol. wosaes «torte see cimoce 1,102,500

a5 GEMSUMAT Yess. n-ne ssc scomeoesooes 1,000,000 -

5 yd yMUny eve ea evesntscs oseeeee 16,400

a COLES Fees score ae 95,750

if SDP. wiaeeae teen veoteete tere 7,500,000

Corethromicriophilumy 72.7... .20:- 00+. see 0s eees 400

INitzsehiasseriata, 2222). tn anescte se se stoustals mene 107,000

Rhizosolenia ‘semispina | -........--ss.0:.--5-e 43,300

.s stolteriothil....es.cc-ssscosceeetee 33,600

Le styliformis ........... eae eee 12,600

Mhalassiosira ‘oraviday -. J... .2: /-s- ese soescere 7,500

oe nordenskioldii .................. 2,500

Peridantumgsppisss.c- eecaes toss cose oetnen eee 10,000
Ceratiumiiusisy. si. -e2o-ascseecnecosseateecee ees 00s

oe ANGERIMEGIUMI 1S... cece sseceeeeeeee 8,000

= [MS ATO fee Cocoa dence ene ace eons 420

a IGN LP OSs coe oe sev wSenctecee vest oece 3,350

33 GEEDOS Sass cots cose res. eon eee 11,350

Pinibinni dae: Gees ccdceace sesso wesceesteseoesceeeee 10,000

Bivadne nordmanni)-5--ee-ee-. senescence 840

Pseudocalanus elongatus .................-0 420

Acartiad iclavist: C2. \ccsuoscess: cc omoeeeee ee 840

Oithona ‘eimuillis Bo <2. s.-ox cee eeee eee are 1,450

Nau p lit 260. alin. ocah ootoasoaegacs saeco newer 6,000

Bera creiecrnrt oe aiecie ore emiaceistee tear 630

Oikopleura

_ This is a typical phytoplankton. It was taken late
in the summer and it is the furthest north of our
gatherings off the mainland of Scotland. The number of
Diatoms is enormous, the gathering is dull green and
dense, and, as the list shows, a few species of Chaetoceras
are represented by nearly fifty millions of individuals.

SEA-FISHERIES LABORATORY. 239

4. Moussa Sound, Shetland, 11/8/11. Surface.

Bacteriastrum delicatulum .................008. 53,000
Chaetoceras atlanticum ..............0.cccceee. 42,800
. | SCTE RL ene atk org apeiin e 32,750

a TUGN = SOAR AA SR ee Aan peor oe 1,562,500
GLE TTC seine eee Loe Ee See ae 201,600

as CORSHMICUUIM, 25... asses csees sass 176,400

- PLIOMDU TINY 2 scatseeceeee envcven sos 1,193,000

b GeMileramee ec ae eee Ssh 5,812,500

a PICRIMICMS mekcusccaracet bens coos ce - 1,250,000

as Rey MIME 8 ss shen hoc acdaen sae 3,625,000

as (ASST tener een ee ihe anne creat nr 438,500

- SEWER ce Seis cia cds aetitiecistawists cates ndais 29,062,500
Maehyesalen TENWIS ° ..2..5...-.cccncnsenceetes 54,800
WG AMAPEANZOCIACUS 2.25 .1chestpenderssesscecenes 45,350
Meprocylmdrus danicus: -............0.c.cese00- 37,800
MEARE MIDUSCTIAUA a scinscc ener cicaedeccescce sects 7,687,500
Nose nia BALA ©... ..c.sescccedccvceccesecoece 118,450
a Sholferroubit-, 2.2 5dds.0skse.s 558s 141,100

eA Spy MEORMIS Ss. cess.tocdcdesaecn cose 25,200
Thalassiosira gravida ...... acre ERR meas, 1,900
» AIOLUCHSITOID UI 4.csc% ees s2ckewe ss 115,900
MEIER SSN 9hecfo wn ov no cceein teed ven tees ved xseree 3,150
Pe MMMNDENITE SO Deacon et oc ds desc y sc cus saeiehsSeeiavets 44,700
Mee RIMPELSIUC ATs Dans 5 csis retiesvndstuceercindecsdess 19,500
Xe TST eA et A 41,600

ie MIIPCEIMEOUIM, 5252. daessneeseotonce cos 103,300

m MERIC EIDTLINIE Da tases enicsees ss sonnei cae eos 171,350

es WOISUD OS: =n ois cass nccenegensbessss Scone. 2,500

af RT CROPOTOSS 28 oa0 5 oat Ge csicteboncens 8,200

% VAS tgs which is cd oaaidecarasiescaharcete 252,000
Calanus helgolandicus ................-sssseeees 10
Pseudocalanus elongatus ..............se0e0 630
PREMIO LTNISUS oso cjusos ca oceedensecccsccacncscds’s 8,800
Cemtropages HAMAS ..........cscecccsssececee 315
TAPIA ENTE: | ooo acdeccsccdcesdedcotececveddaccs 6,900
esis ccd binidnie se'sninvislen anise ss'eh seivive’e 3,150
POI POP COATINT osc eecoucnscccconsecoesns 1,250
PUNE AAPETINCOIUIM 2... .2.ccccecccesesectessccacs 630
ALVES scccscnvocae cs ajencteccccsaveesccs 10
IE oe ve oi cin ecesncnaxcetdsesenactrasesne 1
WTEPACIIA, TCULOVOTSA ..ccncccscscccccveccssresss 5,000
PNM UDOU gos desis sins2a0sdcrectecccescvensdussiss 2,500
MIE ow ricci saw nacediisvscessdcencucsnedos 14,500
i LAG oe oo a 630
Me recat cs atdeivesirssriddeesciesessnacessce 3

This far-north gathering, in the Shetland Isles, in
August, is an undoubted phytoplankton with about fifty
millions of Diatoms. It has also, however, an unusually
large number of Dinoflagellates, and a fair number of
Copepoda and other animals, including 5,000 of the

Pteropod Limacina retroversa.

240 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The main points of difference between this year’s
hauls and those of former years are off Ardmore at the
north end of the Sound of Mull, and m the sea around
the Small Isles. In both places the phytoplankton
formerly present is now replaced by zooplankton. More-
over, round the Small Isles the Diatoms are now mainly
of oceanic type in place of being neritic forms. Other
localities are much the same as in previous years; but in
the new ground, further north, there is more phyto-
plankton than in the seas south of Skye.

During all this time (July 8th to August 23rd) the
plankton at Port Erin, in the Irish Sea, was an undoubted
zooplankton composed chiefly of Copepoda and having
few or no Diatoms.

The facts, such as they are, all seem to support the
suggestion, put forward last year, that the most probable
explanation of the presence of huge masses of Diatoms in
the Scottish Seas in summer is that the phytoplankton

remains longer and passes off more slowly as one goes

further north.

SEA-FISHERIES LABORATORY. 241

Appendix—List of the Physical Observations.*

Date. Time. Locality. Temp. C. Ardometer P17-5°
meuly © 7...... ppm wert Erin Bay .......c0.c-cesdercesoes 14° 1:0262 25-5
;, ene earioeebradda, Head. 2... cc..scooseseese- 12-6°

HO. ane. On Contrary Head .....00..cascecee. 125° 1-026 25-04
Poamern Portpatrick ~ ...3.....jccececelectees 1]-2°
11.30 a.m. N.W. by W. of Corsewall ............ 11-4°
P30 pm tO mules S. of Cantyre. -....c06....5 ES 1-0264 25-24
"2.0 p.m. W. of Mull of Cantyre ............... 11-8° 1-0265 25-42
3.30 p.m. E. by S. of the Mull Lt. House ...... 12-2° 1-0262 25-19
Pa women wy. of Gicha Buoy ............000sses 12-4°
7.0 p.m. Lowlandman Bay, Jura............... 12-2° 1:0256 24-59
is ee 9.15 a.m. 2 miles N. of Lowlandman Bay ... —11-2°
EO UAC... ec ceccsscctscescvcsscse 11:3° 1-0264 25-24
tO CATIA os. cckccecsscmeccsececceses 11-8° 1-0262 25-13
Bremen On Hasdale ........c.c..ccccescleccesenns 11-8° 1-0262 25-13
Pee eA DAY... . 2c nesceccocecssesscecces 13-4°
rereeee 0) a.01. Oban Baiy...........c.ccsceccerserecscres 12-8° 1-0256 24-69
PRUETT WISINOLE 5.25. .20cessssectecccaccesecs 14-8° 1-0242 23-66
Pee meesound Of LOM 2... 6.0.0.....008seecees 15-2° 1-024 23-54
meee bynn Of Morven 5......0..0s<sc0cceeeess 15-4° 1-0234 22-98
Peeper ound Of Mull 9 2... cc. sceceececeeete 13° 1-0254 24-53
G30 pi. Outside Tobermory .«..............0 12-2° 1-0258 24-79
mee Piso. Meee LOPerMOry Bay .....<...6c.esssccceness 13-8°
mara On Ardmore, Mull ...........00..c..00 13-6° 1-026 25:23
Pea mi On Cailliach Point ..............2:.000 13-4° 1-0262 25:39
MMOD SEAT ooo ey .ccesewccsareccecsecaness 15-5° 1-0261 25-69
PET MOE LOT: 6. cece caecevorsceesscnscnesevens 12-3° 1-0264 25-4
aL PET VOUA 2a. essaechavesectsassccncncnst 12-3° 1-0262 25:21
emer Camliach, — ..........cn.0secesessecees 13-2° 1-0264 25-56
429 p.m. 3 miles S.W. Muck Id. .............. 15-2° 1-0263 25-83
Gop. Off Loch Scresort, Rum ............ 15-6° 1-0261 25-71
10.30 p.m. Loch Scavaig, Skye ...............0+ LES 1-0238 23°87
| Ee Serr) LOCH SCAVAIS ........nerscccsscsnccsedss 14-5°
11.30 a.m. Between Canna and Rum ......... 13-5° 1-027 26-2
Peete Eyskeir Id. ........2.0s0ccecceseens 14-6° 1-0267 26-11
4.10 p.m. Between Hyskeir and Barra ...... 15-8° 1-0265 26-14
10.30 p.m. Castle Bay, Barra .................000. 13-9° 1-027 26-28
MR RAs scene Reema, Vatersay Sound ........cccccocccsessses 12-9° 1-027 26:1
12.30 p.m. 10 miles S. of Castle Bay ............ 14-8° 1-0266 26-04
PEMMEPRTO OF CATING oc... cccccssecsscnssncorsce 12-4° 1-027 26-01
fem, och Scresorti, Rum ......ccscecsseeee 13-9° 1-0262 25-49
SLD. s.000 SPE, GC SCLOBOLE. ....25.cce0ccccsesedevcenes 13-6°'
9.30 a.m. Between Rum and Skye ............ 13-1° 1-026 25°14
12.20 p.m. Entering Sleat Sound.................. 12-2° 1-026 24-99
3.0 p.m. Outside Narrows of Skye ............ 12-6° 1-0256 24-66
mereen.tx, Car Crovlin Island <..0.2...0cecescanese 14-2° 1-0254 24-74
BRO, .00. MEI, FOrtred, SKVC .....0..ccovessesceussecers 11-6° 1-0266 25-48
Ra POLUSOD. Ooi ciscnaxsncsvanvates’ecsavsvepere 13-4° 1-026 25-19
1 Be SIT ERIULTOO: scence cetddcvsscancesvsscavetsuance 12° 1-:0266 25:55
wera, OF Holm Island .......0..seccessesses 12:2° 1-0266 25-58
moe p.m. 4:miles N. of 8. Bona ....ccsssecess 12-6° 1-0266 25-65

* As we have stated above (p. 228), we have now reason to think that all our ariiometer
readings are consistently too low, and require a correction to be applied before they can be
compared with other series.

eraecce

eoecee

eeeccce

eeceee

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Time.

8.40 a.m.
10.15 a.m.
11.0 a.m.

6.0 p.m.
10.0 a.m.
11.30 a.m.

LO} pen.

8.30 a.m.

8.50 a.m.

9.10 a.m.
11.30 a.m.

3.45 p.m.

4.45 p.m.

8.40 a.m.

9.30 a.m.

1.0 p.m.

9.0 a.m.
11.30 a.m.

2.40 p.m.

5.30 p.m.

8.30 p.m. -
11.30 a.m.

11.30 a.m.
12-45 p.m.
4.30 p.m.

12 noon

3.0 p.m.
4.0 p.m.
10.0 a.m.
10.30 a.m.
10.50 a.m.
11.30 a.m.
4.0 p.m.
11.30 a.m.
40 p.m.
8.0 a.m.
11.0 a.m.
1.0 p.m.
10.0 a.m.
11.30 a.m.
6.0 p.m.
12 noon

9.0 a.m.
3.0 p.m.
6.0 p.m.
10.0 a.m.
6.0 p.m.
6.30 a.m.
10.0 a.m.
1.0 p.m.
4.0 p.m.
7.0 p.m.

Locality. Temp. C. Ariometer 17-5°
Near mouth of Gairloch ............ 12-8° 1-0252 24-3
Of ua Rea: Ssoscs a-cce eee ce ence 12-6° 1-0264 25-45
Wochwiwey’ s.<vacescaen ase nee 12-4° 1-026 25-02
Entering Loch Torridon ............ 13-2° 1-0256 24-76
Upper Mock, Torridon:..-5---+.eeae 13-8° <1-018 <17-34
Outer Woch* Lormidom= 9... 12-4° 1-026 25-02
Off IN. of Radsay 2.22.27. eee 11-9° 1-026 24-95
iLoch=na=Beisten #2 eacss. 9 eee 12-2° 1-0262 25-19
Off Loch-na-Beiste ...............00. 122 1-0254 24-37
och VAISHM 1225 nch s.r ene eee Ree 12-2° 1-:0248 23:8
Wocl. IDiincleie, eed asee eee cece 15? <1-018 <17-54
och Wig urn © atc sree eee 14-2° 1-0242 23-56
Sound of) Sleatiencececece eee se eee 13-5° 1-026 25-21
Warbets duoch@Nevisiees.ce neces 13:0° 1-0256 24-73
Woch “Newis'ssias-. cedar eae acd). RaS 1-0258. 24-99
Between Eigg and Ardnamurchan 12-8° 1-0262 25:29
Robermory. Bay (avce..eoreeeee 12-9° 1-0246 23-72
Oban w cacs eet eeeereeee feiss 5 12:3° 1-0262 25:21
Olan. 2.0Ute a8 cle eee ere meee 12-5° 10258 24-84
OWA Hacasseeeane eee eee wo atone 12-5° 1:0256 24-65
Oban hal satiate ccsen case eee eee 12-6° 1.0258 24.86
Oil Loch;Spelwer.c.c.csccseeseee eee 11-8° 1-026 24-93
Lynn of Morven and Loch Linnhe, 12° 1-0254 24-37
between Bernera Island and 13° 1-024 23-15
Ru Mor 13-272 1-0238 22-98
Between Sheep Island and Kerrera 12° 1-0264 25:35
Sound of Mull, off Green Islands... 12-9° 1-0255 24-62
N. end of Sd. of Mull, off Ardmore 13-2° 1-026 25-16
S. of Sd. of Sleat, between Higg and ) 13-1° 1-0264 25-54
Sleat Pt., open sea
Isle Onrnsay ie. : ssisecse case ssse sect tists 12-8° 1-026 25-09
Upper: Loch) Mourne. seo: 14° 1-0248 24-11
Outer Moch#riourns e.-sccs-eee eee 14° 1-025 24-31
Sound of (Sleat-2:<.c...ceeseeseeenenens 13-6° 1-0259 25-13
Sound: of Wales \.s5. <5. -secn see eer 13°2° 1-0257 24-86
Woch Alsh=..-cecsaee eee Sopeee ena 12-95° 1-026 25-11
{Off Cro wlimgy cists ssase reece. eee eee 12-35° 10257 24-72
Ott Rui Reai ne etssc tects eee nae 13° 1-0265 25-62
Mouthiot och) Wiwe -nesessese-eecee ee Sy 1-:0251 24-24
Of Handa dsland 222 --cs.e-e eee 12-6° 1-0266 25-65
Woch Wrniloolyy.s.2 sce. caseeess tease 12-6° 1-0267 25°75
@ miles off Tongue /...5.2. 200 cemncesr 13-2° 1-0268 25-95
Halfway, tovHoy 1 -.1-s.s.-0 teased 13-2° 1-0268 25-95
Om, SELOMMNESS $32.55. siweaenceasoeseoeeee 14° 10261 25-4
Risa, Deland: succinic vaste mcdtoneces 13° 1-027 26-11
Scapa Bay. ec: -eacn ssn tices oe emenee 15° 1-0268 26-28
Bring Deeps isa seccn ac. vs deere 3 13° 1-0268 25-91
Ofi Copinsay \cas-ptcadssncedet eee 13° 1-0268 25-91 |
Off S:, of Hairlisland See see.seeet 13:5? 1-027 26:2
Off Sumburgh Head) i22----c..0s-- eee" 14-1° 1-0271 26°41
Moussa Sound. fasssaccer ceeessss-cee 12-9° 1-0273 26-4
IBressaye SOUNG >. .aecuensssceeeeee taser 13-0° 1-0274 26-51
Bressay SOUNG: 2.0.5 eetes-sadeeute eee 12-6° 1-0276 26-63
Of Sumburgh Head y.5...-0-eee eee 141° i-027 26-31
Off Hair-Islandss.etess5.ccer.cssece eer 14-2° 1-0272 26-53
Off IN: -Rotaldsaiy: «.e--ceses- aster 12-4° 1-0272 26-21
Off Nowp! Head igi -ee-csceemecemtet 12-9° 1-0268 25-9
‘
.

—

BE G0 9 Ss SUE Oe bt) Gees a=

wWwwwwwwooc°coecTceo?c
no)

. Off Old Man of Hoy
. Between Hoy and Cape Wrath
. Off Cape Wrath
. Head of Loch Inchard
. Off Handa Island
. Off Ru Coyach
. Off Ullapool
. Upper Loch Broom
. Off Loch Gruinard

. Off Ru Rea
. Kyle of Rona

. Loch-na-Beiste

. N.E. of Rum
. Off Canna
. Off Canna Harbour

. Off Dutchman
. Sound of Jona
. Sound of Iona
. Sound of Iona
. Sound of Iona
. Off Staffa
. Off N. of Mull
. Sound of Mull
. Between Lismore and Kerrera
. Oban Bay
. Off Crinan
. Off Lowlandman Bay
. Between S. of Islay and 8. of Gigha
. Betw. 8. of Islay & Machrihanish.
. Between Cantyre and Rathlin Id....
. Off Garron Pt., Antrim
. Off Carnlouch Bay, N. of Maidens
. Off mouth of Larne Lough
. Larne Lough

. Off Mouth of Belfast Lough
. Off Donaghadee
. On Course 8.8.E. from Mew Id. ...

1 mile N.W.

. 1 mile N.W. Bradda

SEA-FISHERIES

Locality.

eee eee esesseses

SAGO U SICAL. , c.6 cnn co sacedeeemacntae’
Between Rum and Coll .............

oy
Pee eee eee eee eseeesesessesesesese

eee eeeeee
See eee eee eee eee ee eeeeseeee
ey
eeeeee

eee eee ewe ee ese sees eeeee

- (twice)

», (8 miles off Bradda)
», (1 mile off Bradda)
Bradda
Station I, 5 miles out ............+.-
(a mile away)

eee ee es

Station ei Ge

eee eee eee eee eee eee eee ee eee

See eee eee

VL OLOT Handle aaa ear ice aeiics ins roninds’
rN TLR RES RN ee a oe ee
- | LANES PA Shoe ie ae cee

LABORATORY.

Temp. C. Ardometer.

14° 1-0266
14-2° 1-0267
12-9° 1-027
are? 1-0265
12-6° 1-027
13-2° 1-0266
13-2° 1-0263
15° 1-025
13-6° 1-0263
13° 1-0267
13-8°

13-4° 1-0258
15-4° 1-0255
15° 1-0258
15° 1-0264
14° 1-0264
14° 1-0264
14° 1-0262
14-2° 1-0259
14-2° 1-0259
14-2° 1-0257
13-9" 1-0259
14° 1-0258
14-6° 1-026
14-2° 1-0257
15-3° 1-0244
13-5° 1-0258
13-4° 1-026
14-6° 1-0262
15-2° 1-026
13-7° 1-0261
14-4° 1-026
14-3° 1-0259
13-4° 1-0262
13-7° 1-0261
13-4° 1-0261
13-5° 1-0261
14-5° 1-0258
14° 1-026
13-9° 1-0259
14-8° 1-0256
14-4° 1-0258
16-1° 1-0255
15-4° 1-0258
15:7° 1-0257
15-5° 1-026
14-6° 1-0258
14-7° 1-0258
14-6° 1-0259
14-6° 1-0258
14-6° 1-0257
14-7° 1-0258
14-5° 1-0258
14-8° 1-0259
14-9° 1-0259

243

P17.5°

25-89
26-03
26-1

25-59
26-04
25-75
25-46
24-49
25-53
25-81

24-99
25-07
25-29
25-89
25-7

25-7

25°5

25-24
25-24
25-04
25-19
25-1

25-41
25-04
23-96
25-01
25-19
25-61
25-53
25-35
25:37
25-26
25-39
25-35
25°29
25°31
25-19
25:3

25:19
25-05
25°17
25-21
25-37
25-33
25-59
25-21
25-23
25:31
25-21
25°11
25-23
25-19
25-35
25.37

944

Date.

Sept.

?
29°

Sentess
Dace:
TAL as ces
A228
UB nccs os

ae
VOL 6-2.

GS Seis
UD scace

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Time.
Forenoon

4.0 p.m.
Afternoon
Forenoon
Forenoon

Forenoon
Forenoon

Forenoon

Station LI.

pat ey LEE.
Off Niarbyl
Off Dalby

Station I.

oe ee
Station I.
ie We
Off Niarbyl
Station I.
a OU:
Station I.
speae! Lit
yo ILI

Locality.

Coe ceecreseeveseseseneseeseces
Coe coserecececesceseeseeseeses
Cece cerocceroscesscseresnsesce
Coe cceccacescsccecceneseesceses
coerce cceceensecscoseesesseeeee
Ce creccereccsccccesccseceseses
See eeseeeseseesesesecesesevsee

eee eeereesesesesesesecseseoses

15:7°
15-5°
15°

15-2°
14-6°
14-5°
14-6°
14-7°
11-2°
TLS
12°

14-4°
14-5°
14-4°

1-0258
1-026
1-0258
1-026
1-0262
1-0264
1-0262
1-0262
1-0262
1-026
1-026
1-0262
1-026
1-0258

Temp. C. Araometer e 17-53

25-43
25-59
25-29
25-53
25-61
25-79
25-61
25-63
25-03 |
24-93
24-96
25:57
25-39
25-17.
a

SEA-FISHERIES LABORATORY. 245

NOTE ON THE WEST COAST LOBSTER
FISHERIES.

By J. Travis Jenkins, D.Sc., Ph.D.
Superintendent of Sea Fisheries.

In the Annual Report of Proceedings under Acts
relating to Sea Fisheries for the year 1910, issued by the
Board of Agriculture and Fisheries in 1912, there is a
memorandum on the size, sex and condition of Lobsters,
which, as it contains deductions based on erroneous
statistics, needs some correction.

The Memorandum in question has, so far as the
Northumberland district is concerned, been somewhat
severely criticised by Professor Meek. The Board
maintain that it is better to raise the minimum size limit
to nine inches than to protect the berried female. Prof.
Meek points out the fallacy in the Board’s reasoning.*
As a matter of fact, in the Lancashire and Western
District the minimum size limit is nine inches and the
berried lobster is protected, so the arguments on either
side do not concern us much.

The West Coast of England and Wales is admittedly
not an important lobster fishing centre. On this coast
fishing for lobsters and crabs is carried on in open boats,
and the prevailing westerly and south-westerly winds put
a stop to fishing in the winter months. Not only is it
frequently unsafe to venture out to sea in open boats for
days or even weeks at a time, but the lobster ‘‘ pots’ are
frequently washed up on shore or destroyed in other ways
by strong winds and gales.

* Report by Professor Meek on the Memorandum issued by the
Board of Agriculture and Fisheries, on the size, sex and condition of
Lobsters. Printed by order of the Northumberland Fisheries
Committee, 16th April, 1912.

246 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The Board publish the following statistics in the
Memorandum as to the number of lobsters caught on the
West Coast in the years stated :-—

1900... 104,062 1905; ...9 G9taa2
190%, es. 498.826 1906°. 2.2 Gieaaa
1902... 94,948 1907 , .... oe 003
1903... 64,174 1908. 2.2. -SasitG
1904. 3... 485230 1909 ... 46,806

If this table be correct then there is clear evidence
of a rapid and serious decline in the yield of the West
Coast lobster fisheries, a decline which if not checked will
lead to the complete extinction of the fisheries in another
nine years or so.

The Board state :—‘‘ On the West Coast the landings
in the second period of five years are 151,152, or 34°3 per
cent. less than those in the first, and the decrease from
year to year is fairly steady, 1909 showing a decrease of
over 50 per cent. on 1900.”’

‘“There is reason to believe that the statistics have
been fuller and more accurate of late years, so that the
decrease shown ... . notably for the West Coast is
not at all likely to be due to less efficient collection of
statistics.”’

While this last statement may be perfectly true it
does not exclude the possibility of error. The decline
may be due, and as I shall endeavour to show is really
due, to the more efficient collection of statistics and not
to a falling off in the fisheries. The fact is the earlier
totals for the West Coast are unfortunately fictitious
and are far too high.

If this be not the case then it is obvious that the
measures of protection, or the methods of enforcing them,
on the West Coast are not sufficient to preserve the

SEA-FISHERIES LABORATORY. Q47

lobster fisheries from a rapidly approaching extinction.
At the rate of exhaustion shown in the Board’s
Memorandum there will be no lobsters left on the West
Coast in 1920.

That the error in the Board’s statistical returns is a
very serious one may be seen from the study of two
localities for which, fortunately, statistics are available
for the years 1900-9. These two places or ports are
Pwllheli and Holyhead. For ports outside the Lancashire
and Western District the details are not available.

Take Pwllheli first. The total number of lobsters
returned as landed at Pwllheli from 1900 to 1904 are as
follows : —

ii 8100 . 1903 ..: 3.570
1901 ... 3,000 1904 ... 3,692
1902 ... 3,172

As a matter of fact, these lobsters are caught off the
Lleyn promontory of Carnarvonshire, principally at
Bardsea Island, but as they pass through Pwllheli on

** returned ’’ from that

their way to the market they are
port.

As I showed on a former occasion* the records for
Pwllheli previous to the middle of 1902 are undoubtedly
false and consequently misleading. In all probability the
statistics of lobsters for Pwllheli are from 4,500 to 5,000
too high for 1900; that is, there is in one single port an
error of from 4 to 5 per cent. of the total figures for the
whole of the West Coast for that year. Since statistics
are collected from 40 fishing ports on the West Coast, it
will be seen that the possibilities of error are fairly large.

As another example, take the number of lobsters
returned for Holyhead : —

* Lancashire and Western Sea Fisheries, Superintendent’s
Report for the Quarter ending December, 1904. Preston, 1905.

248 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

1900) ey 2,960 19059 22.) Saat
(900 Oke 1906 —... Vee
[902s 25605 1907... vale
1905 fst coo LOS Ree 686
19044 235) hs sts 1909 22% See

Now it is practically certain that the earlier totals
are sheer guesswork. From an intimate knowledge of
the district I have no hesitation in saying that the totals
for 1900 and 1901 are at least twice as high as they
ought to be.

If the official statistics were available in detail it 1s
possible that other similar instances could be quoted.
The above remarks are based on the assumption that the
annual totals of lobsters (in the Board’s memorandum of
1910) have been correctly abstracted from the Annual
Reports furnished by the Central Authority. But have
they ?

A comparison with these annual reports does not
tend to reassure one that such is the case. Up to, and
including 1902, the official fishery statistics were
published by the Board of Trade under the title of
‘“ Statistical Tables and Memorandum relating to the
Sea Fisheries of the United Kingdom.”’

In and since 1903 the Board of Agriculture and
Fisheries have published the official statistics in the
‘“ Annual Report of Proceedings under Acts relating to
the Sea Fisheries.’’

On comparing the totals in the Board’s Memorandum
with these official records, we get the following result :—

SEA-FISHERIES LABORATORY. 249

RETURN OF LozssTERS LANDED (WEST COAST).

Memorandum Official Statistical Tables and
of Board of Statistics. Memorandum relating
Agriculture | (From Annual! to the Sea Fisheries of
os Sie Reports.) the United Kingdom.
1260......... 104,062 155,868 (Board of Trade).
Ci 98,826 160,746
A902.-:.-.0e: 94,943 163,183
HO05..)..... 64,174 138,404
ce 78,237 115,218
iC en 69,572 69,572 Annual Report of Proceed-
i —— 67,997 67,997 ings under Acts relating
ASOT... 51,009 51,009 to the Sea Fisheries.
Tate... 53,706 53,706 (Board of Agriculture and
T9009...) 3. 46,806 46,806 Fisheries. )

According to this there is a considerable difference
in the various official estimates up to and including 1904.
There may, of course, be some explanation of these
discrepancies in the years 1900-4.

The total number of lobsters landed in England and
Wales is also variously given.

RETURN OF LoBsTERS (ENGLAND AND WALES).

Memorandum
of Board of Other Official
. Agriculture Returns
and Fisheries. (as above).
(1910).
E900... 602,346 654,152
POOL, .... 588,571 650,491
; BOOZ. Sass 580,496 648,736
: 1903...... 475,121 549,351
1904...... 515,034 552,015:
) a Ee ee ee aes
‘ 1905, ..5.. 502,673 502,673
‘ 1906...... 520,657 520,657
UL ae 495,326 495,326
1008 2-2... 512,478 512,478
546,805 546,805

_
co
i=)
eo}

As regards the detailed figures furnished by the

4 Lancashire and Western Committee, the Board have

250 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

again fallen into serious errors. For instance, take the
statement (p. [xxxiv) :—

‘““The ratio of non-berried females to males points
unmistakably to the widespread occurrence of stripping.”’

As can easily be proved by a reference to the Board’s
own figures, it points unmistakably to nothing of the sort.

The number of non-berried females and males in the
Lancashire and Western District is given in Table IX.
The next table above is for Cornwall (Table VIII) :—

CoRNWALL. LANCASHIRE & WESTERN.
Non-berried ‘ Non-berried

Females. Males. . Females. Males.

Under 9 in. ...... 986 815 ll 0
oi UO K ter ears 1,061 831 1,643 1,504

ce OME Chess bite ee 723 619 1,386 Iesau

Pee aay. ae 598 442 1,102 @ 1,028
Over’ 2" ee 481 415 759 643

| 3,849 3,122 4,901 — 4,512

That is to say the ratio of non-berried females to
males is in Cornwall 123°2 : 100, and is in Lancashire
and Western 108°6 : 100.

So that if these figures prove that stripping is
prevalent in the Lancashire and Western District, they
also prove that stripping is still more prevalent in the
Cornwall District, and this is very strange since there is
no restriction in Cornwali on the landing of berried
lobsters !

As a matter of fact, the Lancashire and Western
statistics were obtained from two distinct sources, which
should have been kept distinct and not lumped together
as in Table IX. The statistics from Pwllheli were
obtained by our Fishery officer as a result of measure-
ments and determinations of sex of the lobsters landed
by the fishermen. Consequently, lobsters under nine

wa nn aS

SEA-FISHERIES LABORATORY. 251

inches and berried lobsters, the landing of which is
illegal in the District, are excluded. (I find, however,
that the Fishery officer includes 7 berried lobsters landed
on May 27 (2), June 17, June 24 and July 7 (3), 1908,
by a fisherman, who was warned on the first occasion and
prosecuted on the other three.) A few measurements
were made in similar manner by the Fishery officer at
Carnarvon and Bangor.

The statistics at New Quay relate to the fishing
carried on by an individual fisherman (our Fishery officer
at New Quay), who recorded all the lobsters found in his
‘““pots.’’ Consequently, lobsters under nine inches and
berried females are both included. The total number of
lobsters of all kinds caught by this single fisherman will,
of course, bear but a small ratio to those landed and
measured at Pwllheli. At New Quay the berried lobsters
taken in the pots were labelled before liberation, but in
no instance was one recaptured.

To group together returns obtained from entirely
different sources, and from different methods, as was done
by the Board in Table IX is misleading.

The fallacy of the Board’s arguments, which lead
them to conclude there is widespread “‘ stripping ’’ of
berried lobsters in this district, 1s seen when the detailed
statements are examined.

LANCASHIRE AND WESTERN LospstTERS (1907-9).—DETERMINATIONS
oF SEX, ETC.

The Records

From all of one
sources. Fisherman
Berried females ...........00.- 29 ;
Non-berried females ......... 4,901 385
EM i eRe a aaah cick oink eee wad nae 4,512 362
9,442 768

252 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Now the proportion of non-berried females to males
is pretty much the same in the two tables, viz., 108°6 to
100 in the first, and 106°8 to 100 in the second, so that
stripping is to be expected as much in the second instance
as in the first.

But as the figures in the second column were supplied
by our Fishery officer at New Quay who fished himself,
they exclude all possibility of stripping. He was not
allowed to sell the fish he caught, so even the possibility
of a little surreptitious gain or profit through stripping
is excluded. And as the officer was unaware of the
purpose of the returns he made, he cannot be accused of
‘cooking’? them to produce any desired result. |

So much, then, for the ‘“‘ widespread occurrence ’’
of stripping of berried lobsters in the Lancashire and
Western District.

253

L.M.B.C. MEMOIRS.

No. XX. BUCCINUM.
(Tae WHELKE)

BY
moe J. DAKIN, D.sSc., F.L.S:

CLASSIFICATION.

The whelk belongs to that class of the cephalophorous
Mollusea, the GAsTRoPoDA, which includes also the lim-
pets, land snails, and nudibranchs. The group is
characterised by the possession of an asymmetrical body,
a well developed head bearing eyes and tentacles, a foot
for creeping, and a shell consisting of one piece only
(univalve). In some cases the shell is reduced consider-
ably, and it may even disappear completely in the adult
(e.g. Nudibranchiata).

The Gastropoda may be conveniently divided into
the two sub-classes:—STREPTONEURA and HUTHYNEURA.
The first of these groups is defined by the nervous system
being involved in the torsion of the body so that the
visceral loop joining the visceral and pleural ganglia is
twisted into a figure of eight. The morphological] right
side of the loop becomes carried over the alimentary
canal‘ to the topographical left side, and the left
half, under, to the right side. ‘This sub-class is also
named Prosobranchia from the fact that, in most genera,
the gills lie anterior to the heart.

The whelk is a representative of the Streptoneura,
and the common land snail is a type of the other group,
Euthyneura; the exact position of Buccinwm can be seen
in the scheme given on the next page.

254 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Crass GASTROPODA.

Body asymmetrical, well developed head, well
developed foot, usually with flat creeping sole. Shell in
one piece, usually coiled in a spiral, but may be reduced
and completely disappear. Pallial complex situated on
the right or left side in a more or less anterior position.
One half, usually the morphological left, of the a lial
complex reduced, and may be absent.

Sus-cLass I. Streptoneura (=Prosobranchia).
Sexes separate. Visceral loop twisted into figure of
eight. Pallial complex placed anteriorly. In most,
only one gill, which is situated in front of heart.
Chiefly marine. Shell very rarely absent.
Order I. Dotocardia (= Aspidobranchia).
Order II. Monotocardia (=Pectinibranchia).
Heart with one auricle. One gill with
leaflets on one side of an axis. Well
differentiated osphradium. Lye a closed
vesicle. Single kidney. Siphon and Pa
usually present.
Sub-order JI. Architaenioglossa.
Sub-order II. Taenioglossa.
Sub-order 111, Stenoglossa.
Pectinibranchs with much concen-
trated nervous system. Proboscis,
siphon, and penis always present.
Tribe I. Kachiglossa.
Radular formula. I.1.1.
Fam. Buccinidae

Gen. Buccinum.

BUCCINUM. 255

GENERAL DESCRIPTION.

The body of the whelk is divisible into three obvious
external regions, head, foot, and visceral mass (PI. I,
fig. 7). A large part of the animal can be extended
beyond the mouth of the shell, but the visceral mass
always remains hidden, and the entire animal can be
retracted when disturbed. The integument of the
visceral mass is produced to form that characteristic
molluscan structure—the mantle (Pl. I, fig. 7, Pall.).
The mantle forms a continuous cloak round the body, its
free edge being just visible at the shell mouth when the
animal is extended. It encloses a space, the mantle
cavity, which is best developed on the dorsal and anterior
surface. The shell is secreted chiefly by the epithelium
of the mantle, particularly of the mantle edge. The
organs in the mantle cavity will be referred to later.

The head of the whelk in an extended condition
bears anteriorly two appendages, the tentacles (fig. 7,
Tent.). These are compressed dorsoventrally at their
base but are produced to a fine conical lip. They are
capable of considerable extension and contraction, but
cannot be introverted. At the base of the tentacles and
on their outer sides are a pair of cephalic eyes, situated
on slight lateral prominences.*

Below the tentacles and in the middle line is a
conspicuous opening, an apparent mouth. This is, how-
ever, not the true entrance to the buccal cavity. The
latter opens at the extremity of a retractile snout but has
been carried backwards, owing to an ingrowth of integu-
ment, and consequently the true mouth is only seen when

* One specimen of Buccinum undatum found at Port Erin possessed

three tentacles—perfectly normal in shape and each with an eye at
the base. From the position it is probable that a second tentacle and

eye was present on the left side. A similar case in Patella vulgata
has been recorded by Bateson.

256 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the invaginated tube is everted. This eversible tube is
the proboscis, and it can be extended about two inches
outside the opening of the apparent mouth.

The Foot forms the greater part of the mass seen when
the animal is fully extended (Pl. I, fig. 7). It has a very
different appearance in life from that seen even in the
best preserved dead specimens, where the foot is hard and
always somewhat contracted. It is formed of a complex
and powerful mass of muscles, and when fully extended
is perfectly smooth, without any trace of wrinkles, soft
and velvety to the touch, and capable of much change of
shape. It has a perfectly flat ventral surface, with a
blunt anterior and a tapering posterior end (PI. IT, fig. 9).
The ventral surface or sole is used for creeping, but I
have also seen the anterior part used for holding food
matters. When a whelk supports itself above the water-
line in an aquarium tank, it does so solely by means of
suction. Some little force is required to detach it, but
the sole is simply slimy and no adhesive substance is
secreted.

Upon the dorsal but posterior region of the foot is
situated a horny disc, the operculum, used for closing the
aperture of the shell when the animal is withdrawn.

Running transversely across the anterior part of the
foot is the deep pedal groove. This will be described in

detail below, in the section on the foot.

THE SHELL.
The shell of the whelk (Pl. I, figs. 1 and 2), secreted

by the mantle, consists of a single valve which is coiled
spirally owing to the varying conditions under which
different parts of the mantle edge secrete shell
substance. In most cases the twist is of such a
nature that if the shell is held by the apex with
the aperture away from the observer and turned down-

BUCCINUM. 257

wards, the aperture will lie to the right of the columella.
Shells coiled in this manner are ‘“‘ dextral,’’ but one
occasionally finds a ‘“‘ sinistral’? whelk with the spiral
reversed and the asymmetrical viscera developed on the
opposite side of the body.

The shell may be regarded as a long cone coiled into
a spiral. Text-fig. 1 shows the terms in use for the

Apex

Sutures

— Columella

_~ - Outer Lip

Mouth

Anterior
Canal or
Shell Siphon

Frae.. 1.

different parts. The apex is the oldest part of the shell
and often in gastropods presents important characters,
such as being coiled in the reverse direction.

The whorls are in close contact and are about six or

258 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

seven in number in an average adult specimen. The last
turn of the shell is known as the body whorl and is
extremely large. The successive whorls of the shell in the
female show a greater enlargement than is the case in male
shells. The lhnes marking the point of contact of two
successive whorls are known as the sutures. The mouth
of the shell is produced into a short anterior canal or
shell siphon (fig. 1, Pl. I) in which lies a prolongation of
the mantle, the pallial siphon (fig. 7, Siph.). This con-
dition 1s characteristic of carnivorous gastropods. The
pallial siphon can be extended some distance out of the
shelly canal, into which it is retracted when the body is
withdrawn. In some gastropods there is also an anal or
posterior canal which is represented by the perforation in
Fissurella and the series of holes in Haliotis.

The outer surface of the shell 1s covered by a horny
layer which can be stripped off quite easily. In worn
specimens it is frequently absent in patches. This layer,
the periostracum (Pl. I, fig. 4) gives the shell a some-
what brown appearance and a furry surface. It will be
referred to in detail below. The external surface of the
whorls is marked by very distinct grooves and ridges
- which run regularly in a longitudinal direction, and can

be traced round all the whorls to the apex of the shell.

They correspond to the lines radiating from the umbo of
a lamellibranch shell. In Buccinwm these lines are
arranged on crests and troughs; and are in groups of
about six ridges between two crests. The crests become
much more distinct as one passes from the mouth towards
the apex of the shell, where they are more crowded.
Running roughly at right angles to the former
system of longitudinal crests, and therefore transversely
to the direction of growth, is another system. This is
made up of two sets of markings—lines of growth and

bal

BUCCINUM. 259

broad waves or corrugations. The lines of growth are
rather indefinite striae, not nearly so distinct as the ridges
of the first system, except near the mouth of the shell
and particularly on the shell siphon. The corrugations
on the other hand are much more definite on the earlier
whorls, and on the last or mouth whorl of an adult shell
they are only well marked near the suture line.

These transverse corrugations relieve the monotony
of the plane surface and are one of the first characters
which strike the observer when comparing the shell with
that of Fusus.

A longitudinal section taken through the body whorl

_ of the shell shows the following structure :—(a) an outer

wide layer of irregular columns; (6) a middle and
narrower layer, also composed of columns, which are,
however, regular in shape and arranged at almost a right
angle to the surface of the shell; (c) an inner layer
characterised by delicate oblique cross lines.

The outermost layer begins at the outer lips of the
shell mouth, the middle layer commences a little further
inside, and it is soon followed by the inner layer.
According to Tullberg, who seems to have made a careful
study of the structure, a fourth and more internal layer
still, occurs in the older whorls and increases in thickness
as one approaches the apex of the shell. This is not seen
in the micro-photograph, which is from a section through
the wall of the body whorl. Text-fig. 2, after Tullberg,
indicates the position of origin of the layers. —

In addition to these layers of shell substance, there
is a very well developed periostracum which can be quite
easily peeled off from the shell and examined without
sectioning.

The Periostracum is a chitinous layer, yellow in
colour, and raised on the external surface into a number

2960 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of papillae which give the shell the appearance of being
covered with a velvety tissue. These papillae are all

directed backwards towards the apex of the shell. In

sections (Text-fig. 2) the periostracum appears to be
made up of layers lying obliquely to the surface. A
spine is simply a prolongation of one of these layers.

The outer layer (a) of shell (Text-fig. 2, a) is marked
by the possession of more organic substance than the
other layers. The calcareous substance crystallises
irregularly as mentioned above, and the appearance can
be understood better perhaps from the photograph
(Pl. I, fig. 6) than from any description.

The middle layer, Text-fig. 2 (6), is, as we have seen
above, marked in longitudinal sections by parallel lines
running perpendicular to the surface of the shell. In
transverse sections, however, the appearance of this layer

eine Ae, inact ni TAN NR
Lstaiiued ua aie HN ean Ny VOR
Li be

iG 2

is, as Tullberg remarked, just like the inner layer (c) in
longitudinal section. This structure will be understood
better when the remaining two layers (c and d) inside it
are examined in transverse section. It will then be seen
that the layer (d) is marked like the middle layer (6), and
thus both (6) and (d) seem built of parallel columns in
longitudinal sections, whilst they are marked by oblique
lines in transverse section. The layer (c) is apparently
built up of parallel columns in transverse section. The
reason for this strange appearance is that the three inner
layers (b, c, and d) are built up in exactly the same way

SE: Se Oe eS ee ee

BUCCINUM. 261

of small plates which are arranged in rows with the
plates at an angle to each other. Whilst, however, the
rows of the layers (b and d) are situated in a line at right
angles to the direction of the whorls, the other layer has
the rows running almost in the direction of the lines of
growth. It follows that in a longitudinal section of the
shell, the plates of layers (6b and d) are seen trom their
cut faces and appear as columns, the cross striation
marking the cut faces of the plate. The plates of layer
(c) are, however, cut so that they are seen in side view,
and the oblique running lines mark the edges of the
plates. We might therefore divide the shell into two
layers, an outer and an inner, the latter with three
subsidiary strata built up in the same way, but, as the
geologists would say, unconformable. We have already
seen that at the apex of the shell there are a number of
partitions cutting off small chambers. These are formed
entirely by layer (d) of the shell.

sPormation of the shell-layers and
periostracum.—the shell is formed by the entire
surface of the mantle, but chiefly by the mantle edge.
The periostracum and the three outer shell layers are
formed solely by the edge, each of them farther from the
actual margin, whilst the innermost layer (a4) can be
increased in thickness throughout life by the mantle
immediately below it. The structure of the mantle edge,
with the shell secreting cells, will be given in the section
on the mantle.

There is probably little doubt that the actual
crystallisation of the shell substance into the structure
seen in sections takes place outside the secreting cells, and
is determined to a certain extent by the constitution—a
mixture of conchiolin and lime—of the secretion. The
origin of the complex shell structures must, however, be

262 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

further governed by an architecture already present in
the secreting area of the mantle. Such a differentiation
of the secreting cells is, however, ultra-microscopic, and
the sculpture of a mollusc shell belongs to the same
category as the regular shape of the teeth on the radula,
the form of placoid scales and the growth of the Echinus
plates. :

The Columellar muscle is attached to the shell by the
same kind of cells that are noticed in Pecten and other
lamellibranchs. During life a movement of the muscle
takes place, but there is no actual movement of the
muscle fibres. A growth of new fibres takes place in front,
and resorption behind, so that as a result the whole
structure appears to move forwards.

Tue Foor.

The ventral creeping foot is exceedingly well
developed in Buccinum. It is muscular, and capable of
very considerable expansion and contraction, so that
whilst soft and almost translucent when expanded, it
becomes opaque and tough as cartilage and difficult to
deal with in dissections when contracted. The flat and
smooth ventral surface, or sole, has, when the foot is
expanded the shape indicated in fig. 9, Pl. Il. Thus the
anterior margin of the foot is broad, and the width
remains constant for some distance backwards until
towards the posterior end it gradually narrows away to a
point. Running parallel to the anterior margin of the
foot is a deep groove (fig. 9, Pl. II, Ped. gl.), which cuts
off an anterior narrow ridge from the major portion of the
foot; and into this anterior groove open numerous glands.
The molluscan foot is usually well provided with glands,
and these may be divided into (a) scattered gland cells
covering the foot, and (b) specialised compound glands.

BUCCINUM. 2638

The latter class includes the byssus gland of the lamelli-
branchiata. In the gastropoda the specialised glands
may be divided into an anterior foot gland, often opening
anteriorly into a transverse groove, and an unpaired
median gland opening into a cavity situated in the
middle line of the foot. It is very probable, however,
that both these are parts of the same system, and it is
generally believed that one or other is homologous to the
byssus gland of the lamellibranchs. The median
unpaired opening is absent in the whelk, but the anterior
glands are very well developed, and the pedal groove can
be observed quite early in the embryo. The portion of
the foot anterior and dorsal to the pedal groove, as well
as the anterior part of the ventral surface, may be used
as a clasping organ, and in this way the whelk can to a
certain extent retain its hold whilst using the proboscis
and radular apparatus to bore through a molluscan shell
or a crustacean exoskeleton.

As stated above, the foot is highly muscular. It is
almost entirely composed of muscle fibres, and moreover
the greater part of it is one muscle—the columellar
muscle of the shell, which arises from the columella
(Pl. I, fig. 3), and in average-sized specimens from the
inner surface of the 5th whorl, and is inserted into the
under surface of the operculum, and thus must pass
through the foot in order to reach this point. In the foot
it is crescentic in section, the convex side being dorsal
and very distinctly marked off from the narrow band of
more superficial tissue. The whole muscle les here near
to the dorsal surface of the foot. The attachment to the
operculum is on the ventral side and forms an elliptical
area which is situated eccentrically.

The columellar muscle of the gastropods has been
homologised with the adductor of the lamellibranchs by

264 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

those authors who have considered the operculum
homologous with the other valve of the lamellibranch
shell; and with the retractor muscles of the foot, by those
who consider the operculum as representing the byssus
of the lamellibranchs. Both theories are untenable.
The operculum is probably a new structure, and the
search for homologies is sometimes carried too far.

The foot exhibits very great changes in size,
according to whether it is fully expanded or completely
contracted. Whilst the whelk is creeping about, the foot
is almost constantly changing in shape, and if not in total
volume the size of different regions at least varies. This
power of change is due to the vascular supply and the
muscular structure. The pedal arteries form a very
complete network extending throughout the foot. Blood
runs back through a large central sinus. The expansion
takes place through the forcible intrusion of blood into
the foot. If the foot of an expanded animal be suddenly
cut off with a very sharp scalpel, the sudden outflow, or
jet of blood, seen as one severs the sinus, is most striking.
This is due to the muscular contraction which begins
immediately the foot is touched with the knife. The
blood, in fact, has most important functions to perform

in effecting both the expansion of the foot and the

extrusion of the proboscis.

The creeping action of the gastropod foot has already
attracted considerable attention. In most cases when the
animal is in motion a series of waves can be seen coursing
along the foot. These may be in a direction from behind
forwards—direct (Aplysia, Doris, etc.,) or in the reverse
direction anterior to posterior—retrograde (Littorina
littorea and others). In some gastropods again the foot

is divided into two longitudinal halves and these move

alternately, both exhibiting systems of waves. In

od a i | ie

BUCCINUM. 965

addition to the above types there are cases (Parker) where
there seem to be no waves at all and the foot glides like
a Planarian over the substratum. In Buccinum the
waves can only be detected at the edges of the foot, but
in addition to this motion the whole of the anterior part
can be moved forward and then attached whilst the
posterior portion is pulled up toit. This is particularly
well seen when the mollusc is removed from the water.
The actual forward motion of the foot is caused by
muscular contraction pulling forwards all those regions
of the foot temporarily raised,—at least when compared
with the rest of the foot. These regions are the waves,
the crests being for the time the fixed portions.

The Operculum is a disc of chitin with a deposit of
calcium carbonate, placed on the dorsal surface of the
posterior part of the foot (Pl. I, fig. 7, Op., and fig. 5).
It is carried on a slight elevation, the “‘ opercular disc,”’
and when the animal is withdrawn into the shell fits into
the mouth, closing the orifice completely. The operculum
is pulled to with considerable force, for, since the colum-
ellar muscle is attached to this plate and the direction of
pull is almost exactly along the muscle, the whole force
of which the muscle is capable can be exerted. The
amount of lime in the operculum of Buccinum is but
small, and the structure is horny in appearance, lacking
the strength of some other gastropod opercula which may
be thick and extremely hard.

Seen from its superior surface, it is marked by very

_ distinct lines of growth which are arranged concentrically

round an eccentrically placed ‘‘ marginal nucleus.’’ The
attachment to the foot is also eccentric, the oval area
lying quite to one side, the side further from the nucleus
and anterior when the foot is uncontracted. Round the
area of attachment the tissue of the opercular disc form

966 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

a collar which projects for a little distance, sheathing the
unattached marginal regions of the operculum. This
‘“opercular lip ’’ is deepest on the posterior border where
there is a greater width of unattached operculum. When
creeping, the operculum is arranged, as we have seen,
transversely across the foot. The anterior margin in
this position is the internal border, the posterior margin

the external border when the operculum is applied to the
shell mouth.

Hic. 2a.

If the operculum is removed from the foot and the
attached or ventral surface examined, a very different
system of striae will be seen. The operculum is, as a
matter of fact, composed of several layers, and the
markings on the superior and inferior surfaces are
therefore quite independent of one another. The area

of attachment, as already mentioned, is situated towards

the anterior margin, and entirely to one side, therefore,
of the nucleus. This area is marked by about ten bands
(in adult specimens), arranged concentrically, but with
only part of any band showing, the remainder being
outside the area of attachment. Furthermore, these
bands are arranged in an imbricating manner (Text-fig.
2a, 6), each one nearer to the centre overlapping its
more marginal neighbour. The area outside the
region of attachment is covered with a somewhat
glossy horny layer, which hides the concentric bands

a s—-——_ ©

BUCCINUM. 267

as stated above. This glossy ventral layer is broadest
under the posterior margin (Text-fig. 2a, a). The
operculum is composed of three layers, a very delicate
superior layer, a median layer of considerable thickness
which is itself formed of numerous laminae arranged
at an acute angle to the plane of the operculum,
and a third and most ventral layer, the glossy one
described above. The thin superior layer is formed by a
pad of cells situated in a cleft below the anterior lip of
the opercular disc. The middle layer is secreted by the
cells of the disc at the margin of the region of attachment,
and chiefly anteriorly. As a matter of fact, the
epithelium of the foot is perfectly continuous underneath
the operculum, and it is by means of these cells that the
muscle fibres are attached to the operculum. The most
ventral layer of all is produced by the cells of the
opercular lip. It will be noticed that this lip is much
deeper and more obvious altogether posteriorly where the
glossy layer is widest and best developed.

It is possible without more than decalcification to cut
sections through foot and operculum. ‘These will show
quite easily the positions of the various secreting cells.
The secreting cells are characterised by their great depth.
They are narrow and about four times as deep as the other
epithelial cells of the foot in the vicinity.

The pedal groove (PI. II, fig. 9, Ped. gl.; fig. 10,
Ped. gr.) is a deep incision running transversely across
the anterior margin of the foot. It appears quite early
in the larva, and is relatively very large at this period.
In transverse sections of the groove (Pl. II, fig. 10), or
longitudinal sections of the foot, a region round the base
of the groove can be seen with the unaided eye to be
different from the rest of the foot. Sections stained with
methyl-blue-eosin are very characteristic and make

yee

268 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

obvious the fact that this peculiar region is composed of
numerous compound glands.

The groove is, in fact, nothing but a slit-like common
opening of a very large number of glands. The foot in
the region of the groove is bounded, as elsewhere, by a
layer of deep epithelial cells with characteristic striated
cuticular margin and cilia. ‘These ciliated cells are
compressed laterally and separated by gland cells. —
Exactly the same type of cell lines the groove, and there
is no change even at the bottom of it, except that the
cilia are longer and much better developed than else-
where. Here the epithelial cells are much compressed
and the nuclei are drawn out into long spindles.

Between the cells open the compound glands. Below
the epithelium is the muscular tissue, built up mainly of
transverse running fibres, so that their cut ends appear in
section (fig. 10, Ped. mus.). There are, however, scattered
oblique and longitudinal fibres. In the region of the
groove the muscular tissue remains absent, and its place
is taken by the glands. Thus the very distinct demarca-
tion of the gland is due as much to the absence of muscles
here as to the presence of the gland cells. The glandular

tissue (fig. 10, Ped. gl.) is very characteristic. There is

no blue staining contents characteristic of the mucous
gland cells or similar cells in the mantle and pedal
epithelium. Instead, there are packets of very thin-
walled cells situated some considerable distance below the
epithelium.

The packets are bounded by very thin connective
tissue sheaths, but this is so delicate that it may seem no
more than the outer walls of the cells which are arranged
to form the packet.

The cells are intensely vacuolated. From each
packet a narrow path of the same cells runs to the

BUCCINUM. 269

epithelium. These cells, however, do not enclose any
canal. It appears as if the secretion must reach the
groove by passing through the cells. Usually in sections
there are no traces of secretion in the groove, and in living
specimens no mucus or other fluid appears coming from
the groove. What is, then, the function of the gland? It
is homologous with the pedal pore of many other gastro-
poda; once considered an aquiferous pore by which water
entered the animal. The gland secretes the substance of
which the egg capsules are formed. This fact, noted in
1899 by Cunningham to apply to Buccinum and Murex,

has since been found to be true for Purpura by Pelseneer.

Tur MANTLE AND PALLIAL CAVITY.

The pallial cavity proper is the space between the
mantle and the dorsal surface of the body of the animal.
Its floor is formed by the body wall, its roof by the mantle.
It will be advisable to refer in a general way to this part
of the animal in a separate short section, inasmuch as the
cavity contains several important organs belonging to
different systems.

These organs considered together may be termed the
Organs of the Pallial Complex. Three of them are
structures developed largely from the mantle itself—the
ctenidium or gill, the osphradium, and the mucous gland
(Pl. II, fig. 8). Furthermore, there are to be considered
the Rectum and Anal opening (fig. 8, Rect.), the Renal
opening, and the male and female genital openings. The
mantle itself is thick and muscular, and this applies most
markedly to the free edge. The edge is slightly recurved
outwards, and just behind the extreme margin and on the
outer surface is a delicate band of yellow pigment.

If the mantle is slit down the extreme right side on
the left of the rectum (and the oviduct in the female),

te
a

270 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and turned over to the left, the organs of the pallial
cavity can be easily made out. On the extreme left, and
arising from the inner surface of the mantle, are two
ridges which form the side walls of a groove, the pallial
siphon (fig. 8, Siph.). These side walls and the basal
part of the groove are continued so as to form a truncated
cone with a gutter down one side of it. This pallial
siphon lies in the shell siphon, but can be extended a
considerable distance when the animal is active. Water
entering the mantle cavity passes in by means of this tube,
which is characteristic of the carnivorous gastropods.
One would imagine from observation of the living animal
that the siphon was connected with some important system
of sense organs. It is continually in motion from side to
side, and extends much further from the animal and is
more active than the tentacles. | |

Examination of the mantle cavity in this light reveals
an interesting series of organs. The osphradium, which
is a darkly pigmented structure on the left side, is
situated right across the end of the siphon (fig. 8, Osph.).
Thus all water entering the pallial cavity must pass over
it before reaching the other organs.

The osphradium is a narrow organ composed of two
series of leaflets arranged along the sides of a central axis.
It will be discussed further in the chapter on the sense
organs. To the right of the osphradium is the gill
(fig. 8, Ct.). It is separated from the osphradium by the
ctenidial axis which can be seen as a white ridge running
from the most distal part of the gill to the inner end of
the pallial cavity. The gill itself is composed of leaflets
arranged on one side of this axis only, the topographical
right. Between the ctenidium and the cut side of the
mantle the inner surface of the latter is occupied by the
large mucous or Hypobranchial gland (fig. 8, Mu. gl.),

BUCCINUM. 271

which extends therefore from gill to rectum. The
gland is made up of a number of deep lamellar foldings
of the mantle, about twenty in number. The structure
of this organ is considered elsewhere.

The Anus (fig. 8) is situated at the apex of a
prominent papilla on the right side (topographical).

The vaginal portion of the oviduct is conspicuous in
ripe females as an opaque white cylinder on the extreme
right. Its opening into the pallial cavity is not so
prominent as the anal opening by reason of the lowness
of the papilla. In male specimens the pallial cavity will
be filled by the large penis which usually lies twisted
backwards. All these organs terminate about the same
distance from the mantle edge and thus leave free a wide
region, the inner surface of the thickened margin.

The Renal opening is a slit-like pore, situated to the
left of, and slightly above the rectum on the posterior wall
of the mantle cavity, in fact on the membrane separating
this cavity from the renal organ. The long axis of the
sht is dorso-ventral in direction.

The Mantle Edge.

A great part of the mantle, whether at the thickened
edge or in the region of the ctenidium and other organs of
the pallial complex, is composed of a modified connective
tissue. One sees in sections practically nothing but thin
cell walls with nuclei adhering to them, and here and
there fragments of muscle fibres. This characteristic
mantle connective tissue (figs. 31, 2 con. and 45, Pall. gl.)
is seen very well in the thickened edge, where it
occupies about ? of the total thickness. Against the
epithelial layer, which bounds the surface of the mantle,
and underlying this everywhere, is a thick sheet of
compact fibrillar connective tissue of the more normal

272 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

invertebrate type (fig. 45). Running through this
layer are muscle fibres of which the circular are near
to the surface whilst longitudinal fibres occur more
frequently nearer the central connective tissue mass.
The most important feature of the mantle edge is the
epithelium, for this is concerned here with shell building.
The epithelium covering the free inner surface of the
mantle is formed of columnar cells whose height is several
times their breadth. These cells are typical ciliated cells,
so that the epithelium presents here a ciliated surface.
The ciliated cells are separated everywhere by mucous
cells. These ciliated cells extend round the edge of the
mantle to the shell side. On this side of the mantle (and
in that region where the mantle forms the roof of the
pallial cavity) there is a remarkable gland running across
and opening to the surface not far from the mantle edge.
The gland is situated deep below the epithelium imbedded
in connective tissue. The actual gland cells communicate
with the surface by long processes which are so crowded
together that, just below the epithelium and away from
the gland cells, they appear like the fibres of a broad
nerve. Theresemblance is quite striking in methyl-blue-
eosin stained sections, for the stain is not unlike a nerve
stain. Another very striking feature of the gland 1s that,
instead of the fibre-like communication to the surface
opening between epithelial cells, the epithelium appears
to be absent for a short space and its place taken, in fact,
by the gland cell processes themselves.

This glandular mass in the Bucconuwm mantle was
noticed by Tullberg, who discusses its function without,
however, coming to any definite conclusion. He states
that it might very well be a gland for the secretion of the
Periostracum, and this is supported by the fact that the
gland is absent on the mantle below the visceral mass

,
a
.

BUCCINUM. 273

where no Periostracum is formed. Against this, however,
he adds that the gland is too large for this function alone
and that it would be peculiar to find a special gland for
the secretion of the Periostracum, whilst the shell itself
is formed by the general epithelium of the mantle.

In my opinion the objections that Tullberg brought
forward are not important. In the lamellibranchs the
Periostracum arises in a groove from a very definite
pad of cells, certainly epithelial in position but still
differentiated enough to form a special organ. Hence
there is no reason why the thick Periostracum of the
whelk should not be formed by this gland. In any case
no other function has been ascribed to it. The compara-
tive anatomy of this organ is being followed up by the
author.

The shell side of the mantle from the opening of the
gland inwards is faced by epithelial cells differing from
those already noted in the absence of cilia. These are the
shell secreting cells. They are marked, particularly near
the gland, by the possession of granules of yellow
pigment.

RESPIRATORY ORGAN (CTENIDIUM).

There is only one ctenidium present in Buccinum, as
in most of the higher Gastropoda. This is the morpho-
logical right gill, but is situated now on the left side of
the pallial cavity. It has already been referred to as
being visible through the thickness of the mantle. This
ctenidium extends from a point, in line with the anterior
limits of the osphradium and mucous gland, as far back
as the pericardium (fig. 8, Ct.). It is composed of a large
number of flattened leaflets which are packed parallel to
one another and vary in size, so that they become

274 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

successively smaller as either end of the ctenidium is
approached.

These leaflets are roughly triangular in shape. The
axis of the ctenidium lies close to the osphradium (fig. 8,
Ct. av.). The respiratory leaflets are disposed along one
side only and are further attached by the whole of their
bases to the mantle (see Text-fig. 3, p. 277).

The ctenidium is therefore Monopectinate.

The efferent branchial vessel runs along under the
ctenidial axis and thus the area of each leaflet adjoining
the axis may be known as the efferent end and the free
side, the efferent margin of each leaflet. Branchial
lacunae extend up the afferent margins of the leaflets from
the afferent branchial sinus which lies in the mantle
immediately below the point of attachment of the afferent
edges of the lamellae. |

The ctenidial axis (fig. 8, Ct. az.) 1s conspicuous as a
smooth ridge running along the inner surface of the
mantle as far as there are ctenidial leaflets.

It is made of the same peculiarly vacuolated con-
nective tissue (fig. 39, Ct. gl.) seen in the mantle at the
base of the ctenidial leaflets. Towards the side to which
the branchial lamellae are attached the place of this
tissue is taken by longitudinal muscle fibres (fig. 39,
Ct. mus.). This layer increases in thickness towards the
middle of the ridge. |

The ctenidial nerve (fig. 89, Ct. n.) runs along the
axis not far from the osphradial side, and: gives off
branches at intervals, which pass to the leaflets.

HISTOLOGY.

The microscopic structure of the ctenidial leaflets

is interesting by reason of the histological differentiation
of the different areas. Each leaflet consists of a double

BUCCINUM. 975

bounding layer of epithelium enclosing a flattened cavity,
which is a blood space. Underlying this epithelial layer
(figs. 31, 32, 33, ct. e.', ct. e.”, ct. e.’") there is an internal
connective tissue layer with muscles, lining the blood
space just mentioned. Between these two layers a
supporting tissue is developed which is much thickened
near the ctenidial axis and runs along the efferent margin
of each leaflet (figs. 31, 32, 33, Sup. m.). -

This supporting tissue, which in its thickened parts
is apparently hyaline, has been regarded as cartilaginous
or chitinous; this will be referred to below.

The relation of the various structures enumerated |
can be made out best in a section transverse to the leaflets
and perpendicular to the mantle. Such a section, running
through the efferent margin, is figured on Pl. IV (figs.
31, 32, 33). |

The external epithelium, which bounds the lamellae,
differs considerably in the various regions. Taking the
section through a point near the efferent margin, there is
first the free edge to be considered. The epithelium here
is composed of somewhat deep and regular columnal
cells bearing cilia (fig. 33, ct. e.'). This epithelium
diminishes in thickness and becomes more irregular than
indicated in the figure as one leaves the free margin.
The cilia are also lost and the cells appear glandular
(fig. 33.) Following on this region the cells again become
more deep, more regular and with deeply staining cyto-
plasm. ‘They all bear well developed cilia, so that the
whole area occupied (fig. 32, Ct. e.'") by them is very
conspicuous by reason of the marked contrast with the
glandular cells lacking cilia on either side of it. Passing
this area towards the mantle, the cells become once more
glandular and without cilia. The epithelium here is
often thrown into folds through contraction, probably on

| Bi rr

276 YRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

fixation (fig. 31, Ct. e.”), but this in all probability can
take place in life since there is a plentiful supply of
muscles in the sub-epithelial layers.

Immediately wader the epithelium there is a
supporting membrane. This is very delicate and almost
indistinguishable under the greater part of the area of the
leaflets, but thickens considerably, so that it becomes
the outstanding feature in stained sections, near the
efferent margin (fig. 33, Sup. m.). It is this substance
that has been termed cartilage or chitin. It is clear,
however, that this supporting membrane must be con-
sidered in conjunction with the connective tissue on its
internal face, that is, bounding the blood cavity. This
layer is distinctly peculiar. Seen in transverse sections,
all that can be observed are a few nuclei with very little
surrounding cytoplasm (fig. 32, Ct. con.). If, however,
a section is cut very slightly tangential to the leaflet, it
may be possible to secure the layer itself. It then appears
made up of squamous cells, often with the corners drawn
out (fig. 34). If these cells touched, making up a definite
membrane, it would be difficult to distinguish them from
an endothelium. Instead of this, they are more or less
scattered, and at odd places there is a crowding together
to form islands of pseudo membrane. ‘This tissue is thus
very characteristic. Now the supporting membrane is in
close contact with these cells, and moreover, in places it
is quite continuous with the matrix surrounding
them. This brings out strongly its resemblance to the
matrix of ordinary connective tissue both in structure and
in staining. Hence I have come to the conclusion that
the supporting structure, thickened at the efferent
margin, is really a connective tissue, free from cells or
fibres, and that the cells which have formed it occur on

its inner surface.

BUCCINUM. OT

The relative areas covered by the different structures
are indicated in Text-fig. 3.

The connective tissue supporting skeleton extends
from the axis along the efferent border to the angle. At
first it extends from the free edge of each leaflet to the
mantle, but it gradually becomes reduced. The area
of ciliated cells increases quickly and then extends to the
angle of efferent and afferent edges, occupying in sections
from about a third to one-sixth of the length of the leaflet
(measured from efferent margin to mantle side). The
glandular cell area is nearest to the mantle. It increases
steadily in thickness as the afferent edge of the leaflet is
approached and is widest in that region.

at *
gz iN
2

: Li Se
Fic. 3. ct. e.’ Area of Glandular Cells. ct.e.” Area of Ciliated Cells.
These differ from the reference letters on Plate IV.

One further detail remains to be described. Peculiar
characteristic bridges run across the blood cavity in the
leaflet. Each of these appears to be formed of muscle
fibrils which diverge at their extremities (fig. 34,
Ct. mus.). The cell in which these muscular fibrillae
have been formed remains, and is usually quite obvious
with its residual cytoplasm and nucleus in the centre of
the bridge.

Thus it is possible by contraction of these numerous
muscle strands to approximate the two surfaces of the
leaflets, and hence to force out the contained blood.

278 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Another peculiar histological structure may be
referred to here. The epithelium of the lamellae is
simply the folded epithelium of the inner surface of the
mantle. The outer epithelium bounding the mantle exter-
nally is, of course, not folded here or elsewhere (fig. 31,
Pall. ep.). Between these two layers there is the very
characteristic median stratum of connective tissue made
up of extremely large cells with delicate walls, feebly
staining nuclei and practically no contents (fig. 31,
X.con.). What function this layer may play has so far
not suggested itself.

MUCOUS GLAND.

The mucous gland is the most conspicuous organ in
the pallial cavity, both by reason of its secretion as well
as by its structure. It is a modified region of the mantle
between the ctenidium and the rectum where gland cells
predominate, and the inner wall of the mantle is thrown
into deep folds which run transversely, overlapping
slightly and hanging down into the pallial cavity.

The mucous gland is-really the inner wall of the
mantle whose cells are elongated and specialised as gland
cells. The anterior folds of the gland are directed
backwards, the most posterior ones forward (fig. 8,
Mu. gl.). They are much thicker than the ctenidial
leaflets and are separated by much greater spaces. The
number varies, 20-25 being about an average for a full-
sized whelk. |

This region of the mantle is extremely well
supplied with blood, as a glance at fig. 35 will
show, and numerous vessels run down parallel to
the folds from the reno-mucous vessel to the afferent
branchial vessel. The secretion of the gland is either
perfectly hyaline or yellow-white in colour. It 1s
perfectly abominable to handle, and after months in

BUCCINUM. 279

five per cent. formalin it still retains its fresh appearance
and consistency. It can be drawn out into long threads
of surprising length (some feet) without breaking. The
secretion of this matter takes place very rapidly when
the animal is severely stimulated, particularly with
irritating fluids. Concerning its function but little is
known. It would appear in the first place to be a pro-
tective—or defensive—secretion. It does not leave the
animal gradually after secretion but is produced, as we
have seen, spasmodically and quickly, and raises itself
in a sheet. Obviously it removes in this way any dirt,
sand grains or other matter from the organs of the pallial
cavity, and, moreover, protects them to a certain extent
from the entrance of such materials.

HistoLtocy or Mucous GLanp.

Sections taken through the mucous gland show that
we have to deal with a very much modified epithelial
layer, which rests on a basement membrane of connective
tissue overlying the peculiar cellular connective tissue of
the mantle (fig. 40, Con.). This latter tissue extends
into the folds, but only occurs as bridges running across
at intervals and leaving great cavities between the two
sheets of epithelium. The epithelium is composed of
three types of cells as described by Bernard: 1, Mucous
cells; 2, ciliated cells; 3, neuro-epithelial cells.

In sections, the characteristic appearance is to find
cell walls running from the periphery to the basement
membrane dividing the whole up into large chambers
filled with mucus (fig. 40, Mw. c.), but in addition there
is a more protoplasmic peripheral region bearing cilia
and another series of cell walls. There are also two
distinct nuclear regions, one of which is peripheral

280 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

(nuclei belonging to the ciliated cells), and ue omnes
basal (nuclei of the mucous cells).

The ciliated cells are long and narrow, with,
however, an expansion at the periphery which forms a
kind of platform. The mucous cells are very large and
wide, their length varies according to the amount of
mucus present. Their peripheral ends are, however, often
quite attenuated.

Bernard has made a very detailed study of the process
of mucus formation, examining pieces of living tissue
from the gland with the microscope. He concludes that
before the production of mucus the ciliated cells are all in
contact and form a continuous surface without grooves or
openings. The mucous cells do not reach at first the level
of the surface, but gradually they extend until they
appear between the ciliated cells. A small opening occurs
and from it the excretion pours out as a drop. The cell
continues to secrete mucus.

The ciliated cells can detach their peripheral
portions, which go off as ciliated spherules minus nucleus
and with little protoplasm. Finally dead cells of both
kinds are expelled and may be seen in the excretion.

THE ALIMENTARY CANAL.

The alimentary canal opens at the true mouth, at
the apex of a long retractile proboscis (fig. 11, Prob.),
probably not to be seen without dissection in the preserved
specimens. The opening which has been noted on the
surface of the head below the tentacles is not then the true
mouth. At this point the body wall is turned in to form a
permanent introvert of considerable length, part of
which, however, is again turned on itself to form

The Proboscis. Text-fig. 4 explains this part of the

a ae

BUCCINUM. 281

body better than any description. The invaginated body
wall between a and d forms both the proboscis, which
can be protruded, and a proboscis sheath; part of the
latter, however, can be everted (fig. b to c).

a Od

Fia. 4.

The section of the proboscis sheath a-b in Text-fig. 4
is connected somewhat closely to the body wall by short
muscles. This region retains its position whatever be the
disposition of the proboscis. A definite ring of muscle
and connective tissue (usually of a reddish hue in fresh
specimens) encircles the sheath at b, and marks the
boundary of the next section b-c, This portion, about

282 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

13 inch long in the adult, is thin walled, made up of
muscular fibres and connective tissue, and is connected by
long strands of muscle with the walls of the body cavity.
Some.of these strands pass backwards for some distance.
Now this region can be everted in such a way that whilst
forming part of the proboscis sheath in the retracted
condition (fig. 4), it forms the base of the proboscis
in the extended position (fig. B). Thus the protrusion
of the proboscis is not due to muscular action of this
structure itself but to the unfolding of the hinder part of
its sheath. The proboscis is nothing but a further con-
tinuation of the body wall, and the true mouth lies as
mentioned above at its distal end (figs. a and B, M.).

The cavity (part of the external world) between the
proboscis and its sheath (Text-fig. 4, Khyn., and Pl. II,
fig. 11, Ahyn.) is known as the Rhynchodaeum, and
the opening of the latter, or the false mouth, is the
Rhynchostome (/¢st.). Such a proboscis is termed a
pleurembolic proboscis (Lankester*), from the fact that
when withdrawn it is the base that is pulled and
disappears first. The other and opposite type is the
pleurekbolic, met with in the Cypraeidae, etc. The
proboscis of Buccinum was known to the ancients, and
both Aristotlet and Pliny? refer to it. Cuvier was the
first, however (1817)§, who described it with ce and
detail.

Pharynx.—The mouth (Text-fig. 4, W/., and fig. 11)
opens into a muscular pharynx (PI. II, fig. 12, Ph.), the
walls of which are attached all round to the proboscis
walls by radiating muscles. Into the floor of the

* Lankester. Art. Mollusca, Encyclop. Britannica, 9th edit.,
Vol. XVI, 1883:

+ Aristoteles. De Animal. hist., Lib. IV, cap. 4, §§ 7, 8, 9.

{ Phnius. dist.Nat. i. Xie. 87,

§ Cuvier. Mém. pour servir 4 l’hist. et a Vanatomie des
Mollusques.

BUCCINUM. . 283

pharynx, which bears no teeth, projects the tongue
apparatus, and the muscles of this characteristic organ
almost surround the anterior part of the alimentary
canal. The whole structure is known as the Odontophore
(Text-fig. 4, Od.), and consists essentially of a tongue,
projecting into the pharyngeal space, covered by a
flexible rasp—a membrane set with teeth—known as the
radula. A description of this apparatus is given
separately below.

The pharynx extends backwards for about 1 cm., and
almost at the point where the radular apparatus opens into
it, two salivary ducts enter laterally, one on either side
(Pl. II, fig. 12, Sal. d.). These ducts can be seen as two
delicate white tubes running along the sides of the
oesophagus (fig. 12, Oes.), the next portion of the gut.

The Oesophagus extends from the pharynx to the
stomach, and is the longest section of the alimentary
canal. The most distal part, immediately behind the
pharynx, is flattened dorsoventrally and runs along
through the proboscis to its posterior end. Here it turns
abruptly on itself and runs forward again in close contact
with the proboscis sheath. The anterior direction is kept
until the region is reached where the nerve collar lies
hidden by the conspicuous salivary glands, and then
another somewhat abrupt bend occurs (Text-fig. 4,4 and B)
and the oesophagus passes through the nerve ring and
runs posteriorly along the floor of the body cavity. This
curious looping of the oesophagus is probably due to the
nerve collar which has retained its normal anterior
position and compelled the alimentary canal to take the
course which has been described. The part of the gut
which is thus bent into an § is marked by longitudinal
folds projecting into the lumen. None of these
longitudinal folds are specialised or better developed

ae be

284 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

than others. At its sides run the salivary gland
ducts already mentioned. They arise in the large
salivary glands, compact bodies of a yellow colour,
which are situated asymmetrically about the alimentary
canal and nerve collar. The latter is hidden com-
pletely except from below. ‘The left gland les more
posterior and more dorsal than the right, and thus covers
part of the latter. The ducts are formed by numerous
small branches joining up in the tissue of the glands
and leaving them dorsally (PI. II, fig. 12, Sal. gl.). The
histological structure of this organ is given below.

Neither gland alters its position during the evagina-
tion or invagination of the proboscis, and a further most
important point to notice is that nezther salivary duct
passes through the nerve ring.

This is probably due to the fact that with elongation
of the proboscis the salivary glands came to he in front
of the nerve collar. With the later elongation of the
salivary ducts the salivary glands came to lie behind the
nerve collar and moved to the position externally to the
collar. This explains the fact that in the Rachiglossa
the ducts do not penetrate the nerve collar, a feature
otherwise common in the Monotocardia, where the glands
are posterior to the nerve ring.

A little distance behind the nerve collar a narrow
tube opens into the oesophagus on its dorsal surface
(fig. 12). After running forwards for a few millimetres,
it curves round and widens into a very thin-walled bag,
usually flattened, and of a brown colour in fresh
specimens, owing to the contents. The bag extends back ©
for some distance, lying upon the oesophagus. It is the
so-called gland of Leiblein (Pl. II, fig. 12, Zn. g.)—the
poison gland of the Toxiglossa.

Both gland and oesophagus move over towards the

BUCCINUM. 985

left side as the visceral mass is reached. Just before the
region of the pericardium and digestive gland is entered,
a peculiar caecum is to be found on the oesophagus
(fig. 12, Im. g.). Its walls are rather more thick than
the part described above, and resemble the short piece
now left between caecum and stomach. The caecum can
be easily seen in simple dissections, for the oesophagus
has now left the head region and is visible without any
incision whatever at the surface of the visceral mass on
the left ventral edge. The caecum itself lies just behind
the pericardium (fig. 12, Caec.).

The Stomach—that part of the alimentary canal
into which the ducts of the large digestive glands (the
so-called liver) open—is a bag of considerable size, with
one surface lying against the digestive gland and the
other surface against the bounding integument. Thus
the stomach is clearly visible without any dissection.
Curiously enough, it is attached strongly by short muscle
- strands to the epithelial layer of the body wall, so that
the latter, which can be easily removed from the other
parts of the viscera, is only pulled away with difficulty
from this area.

The stomach (fig. 12, St.) is very irregular in shape.
It is flattened, so that there are two more important
surfaces, and it is elongated in an antero-posterior
direction. The oesophagus opens into it ventrally about
midway between the point of origin of the rectum and
the posterior apex of the stomach. Just before entering
the stomach the oesophagus passes under a somewhat
conspicuous lobe of this organ, which is marked with
radiating striae and lies between oesophagus and intestine
(fig. 13, Dg.”).

The markings on the external surface of the stomach
correspond to ridges which occur on the inner surface and

286 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

project into the lumen. Two openings into the stomach,
both on the inner surface, mark the entrance of the ducts
from the digestive gland (fig. 13, Dg. d.). One of these
is situated anteriorly close to the proximal part of the
rectum, the other is just posterior to the point of a
of the oesophagus.

The Digestive Cland is the ee structure in the
visceral mass, and extends from the pericardiac region to
the tip of the spire. It is brown or brown-green in colour,
and soft and oily in consistency, with no very pleasant
smell. This large gland, which is made up of fine
branching tubules, was formerly regarded as a “‘ liver.”
It is now agreed that this name is unsuitable, as the
digestive functions are more comprehensive and unite the
functions of the different digestive glands of the verte-
brate gut. It is, furthermore, the chief organ in the body
for absorption of digested food.

Originally the digestive gland of the Gastropoda was
paired and symmetrical. In the adult Buccinum there is -
an obvious division into right anterior and left posterior
lobes. The latter is much the larger of the two and
extends from the tip of the stomach to the end of the
spire. The boundary of the two regions lies at the
posterior end of the renal organ. The paired nature is

still further indicated by the fact that the tubules, of

which the gland is composed, open into one another, and
finally form two large ducts which enter the stomach, as
mentioned above. The posterior part of the digestive
gland is partially covered by the gonad which lies,
forming a kind of arch (fig. 58), on its dorsal surface.
The Intestine (fig. 12, Rect.), the original posterior
portion of the gut, is of shorter length than the oeso-
phagus. Owing to the torsion which has taken place in
development it runs forward dorsally to open into the

BUCCINUM. 287

pallial cavity. The intestine leaves the stomach dorsally
and anteriorly and lies close to the surface until the renal
organ is reached. It then plunges underneath the latter
(though still on the surface of the digestive gland and
outside the pericardium), and reaches the pallial cavity.

It is now some distance away from the oesophagus
and ascends into the wall of the pallial cavity, taking a
course along the right side at the extreme edge. In the
female this distal portion, the rectum, is compressed by
the oviduct. The rectum terminates in a conical protu-
berance at the end of which the anus is situated (fig. 8,
An.). With the exception of glandular walls, no special
anal or rectal glands are present.

Histology of the Alimentary Canal
and related Organs.

A detailed account of the histology of the whelk’s
tissues would be far beyond the limits of this Memoir.
Only some of the more typical and important structures
will be referred to here.*

The Pharynz.—The pharynx in life has a sean
pink colour, due to its muscular wall. In sections one
finds the lumen of the gut lined by a layer of deep
epithelial cells. As a matter of fact, the lining of the
gut is very similar throughout its length, and the same
kinds of cells are found in the epithelium, viz. :—
(1) Ciliated cells, (2) Eosinophilous cells, (3) Gland cells.
Whatever may be the function of the two latter types, it
is interesting to notice that they occur throughout the
whole length of the gut from pharynx to rectum.

The ciliated cells (fig. 24, Cl. c.) are typical tall

* The author intends to publish eave a series of papers dealing
with etinacan Histology.

+ 2.
;

, 4
iP

288 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

columnar epithelial cells, with an oval nucleus situated
near the middle of the length. A distinct border, due to
the desmochondria at the base of the cilia, is present.
With either methyl-blue-eosin, or Mallory’s stain, the
contents of the eosinophilous cells (small spherical
granules) (fig. 24, Hv. c.) stain an intense red and the
contents of the gland cells ight blue (fig. 24, Gl. c.).

The most interesting part of the pharynx is, however,
its muscular sheath, which underlies the epithelium. It
may be divided into two layers, longitudinal fibres
situated next to the epithelium, and an outer circular
muscle layer (fig. 24, Mus. long. and Mus. circ.). ‘There
is no outer layer of epithelium bounding the cavity in
which the pharynx lies, for this is simply a haemocoele.
The muscle fibres are extremely thick, and each is
surrounded by a capsule of connective tissue. This
matrix is, however, so reduced that it has rather the
appearance of very thick cell walls, where the muscle
fibres are cut transversely. The fibres themselves
resemble the cell contents, for they are almost round in
transverse section; the angular shape really possessed by
them is probably due to crowding and pressure. The
structure of the fibres is extremely distinct in this
pharyngeal musculature, and one sees a_ beautiful
peripheral arrangement of fibrils surrounding a large
central granular sarc, in which may sometimes be seen
the nucleus. A better or more easily procured example
of this type of muscle fibre could hardly be imagined.
These large fibres with their great sarcoplasmic centres
give the reddish pink tinge to the pharynx, a colour which
is hardly ever present in molluscan muscles, and in the
whelk in one other place only, the muscles of the
odontophore.

The Gland of Leiblein.—So far as I am aware, no

BUCCINUM, 289

description of the structure and chemistry of this gland
exists. In life the gland possesses brownish contents
which can be seen through the delicate walls. It 1s
homologous with the so-called poison gland of the Toxi-
glossa, and, as stated above, belongs normally to the
ventral side of the oesophagus. Its opening has been
brought to the present dorsal position by the torsion of
the alimentary canal in this region.

The walls of the sac are formed of a delicate layer
of connective tissue (fig. 30, Con. t.), which is, at the
same time, a supporting membrane for a stratum of
peculiar cells which line the cavity. These are extremely
long pear-shaped cells which are attached to the basement
membrane by their narrow ends. They have the appear-
ance of loosely adhering cells, sometimes looking like
contracted Infusoria, attached by narrow stalks and
protruding swollen sac-like portions into the lumen of the
gland (fig. 30, Gl. c.).

_ The cells themselves are of all sizes, and vary from
ordinary columnar epithelial cells to the elongated pear-
shaped kind. There is no doubt that all the cells are of
one category, and the differences observed are merely
those of growth. In all cases the nuclei, which are
elliptical in shape, are to be found near the bases of the
cells. The cells are well filled with protoplasm and
crowded with brownish yellow granules. In the sections
so far examined the lumen of the gland has contained
numerous cell remains, and it is evident that dehiscence
of the whole cell, or at least part, takes place when filled
with the brown granules.

The function of this gland is at present problematical,
and I do not know on what evidence the term poison
gland, as applied to the homologous structure in the
Toxiglossa, has been given. It could hardly function as

290 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

a poison gland in the whelk, opening, as it does, so far
back. It is in all probability a digestive gland, but it
might function as an “antiseptic.”

The Salivary Glands possess an extremely compact
structure. They become brittle in preserved specimens,
and are most difficult to cut when embedded in paraffin.

Sections show that after the salivary ducts break up
in the gland, the branches dividing into finer tubules in
their turn soon end through complete occlusion of their
cavities. Sections of the ducts with their columnar cells
bearing cilia (fig. 26, Sal. d.) are met at irregular
intervals, but the greater part of the tissue is composed
of groups of large intensely vacuolated cells (fig. 26,
S. gl. c.). These large cells become continuous with the
epithelium of the ducts, but, unlike the gastric gland —
cells, they are so large that the cavity of the ductule soon
ceases to exist when the walls are formed of them. In
most of the cells a trace of protoplasmic contents at least
remains, and there is a distinct nucleus. The rest of the
cell is either one large vacuole with colourless non-
staining contents, or is filled with bodies of irregular
shape and very variable size, which stain intensely with
eosin (fig. 26, S. gl. c.'). ti

In some gastropods the secretion of the salivary
glands contains a large amount of free sulphuric acid,
which is supposed to aid in the boring of calcareous
shells. No marked acid reaction could be obtained with
the whelk secretion. Griffiths finds that the gland has
the same function as that of Patella, and contains a
ferment capable of transforming starch into glucose.

The Oesophagus.—The section figured has been cut
not far from the caecum (PI. III, fig 25). It has already
been mentioned that longitudinal ridges run along the
lumen of the oesophagus. These can often be seen from

a
:
3

BUCCINUM, 291

the exterior, but not because the external surface 1s
thrown into folds. It is only the internal layers that are
folded, so that as a result the walls are alternately thick
and thin. There is no external epithelial layer bounding
the alimentary canal, for it runs through a haemocoele
and not a true body cavity bounded by an endothelial
layer. One finds, therefore, most externally, an attenuated
connective tissue layer (fig. 25, Oe. con. )- This gives
place to compact dense connective tissue, through which
run many muscle fibres (fig. 25, Oe. con.'). Externally,
the circular or transverse muscle fibres predominate,
internally one finds longitudinal fibres, and this layer is
particularly thick and forms the main substance of the
folds. The connective tissue sheath is divided, therefore,
into two distinct layers by reason of the muscles which
run through it. The inner layer with the Iongitudinal
muscles varies in thickness according as to whether
it is under a groove or a fold, and forms the support for
the epithelium which lines the gut.

The Epithelium is composed of regular columnar
epithelial cells, the height of which is about twelve or
more times the thickness. There are three kinds evident,
viz. :—(1) Ciliated cells, (2) Hosinophilous cells, and (3)
Gland cells, and their frequency is in the order given
above, the gland cells being least numerous.

The ciliated cells are very narrow basally, but
increase in thickness towards the lumen, and their
surfaces form a distinct unbroken plane. They are
typical ciliated cells and show very distinctly the double
row of desmochondria at the base of the cilia, and the
connecting fibres in the cytoplasm.

Between these cells occur the eosinophilous cells in
great numbers. They are more common here than any-
where else, and are crowded with small granules which

292 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

show a most marked avidity for eosin. The gland cells
stand out very distinctly in methyl-blue-eosin stained
preparations, for their contents appear light blue, whilst
the epithelium generally is a dense red, a granular red!

They do not occur in very great numbers. In some
sections the lumen of the oesophagus is filled here with
the blue stained contents of these gland cells and the
red stained extruded cells. Hosinophilous granules also
abound in the secretion.

The Caecum.—The caecum differs from the seme just
described in having the simple folds of epithelium and
connective tissue thrown into most complex secondary
folds. Thus the lumen is split up and reduced in size,
and the connective tissue is reduced to a thin layer
between the folds of epithelium.

Another important and obvious distinction is that
the eosinophilous cells have become much reduced in
number, and hence the lining epithelium has almost lost
the granular densely stained appearance. A few
scattered gland cells are to be found. What the function
of this caecum may be it is difficult to say, for the walls
are not in the least like the walls of a gland, and gland
cells are reduced in number. ~

The Stomach.—In structure the wall of the stomach
does not differ essentially from the rest of the alimentary
canal. The cavity is lined by a layer of columnar
ciliated epithelial cells (hexagonal in transverse section),
which are perhaps not so long nor so slender as the cells
of the rectum, but resemble them closely. Amongst
these cells are scattered eosinophilous cells of the same
character as those found elsewhere in the walls of the gut.
Gland cells occur but rarely. The nuclei of the epithelial
cells are to be found nearer to the basal membrane than
those of the cells of the remaining parts of the digestive

BUCCINUM, 293

tract. Underlying the epithelium is a thin but distinct
basal membrane, and below this a layer of connective
tissue with muscle fibres. |

Numerous large blood spaces occur in the connective
tissue, so that 1t may practically be said that the stomach
lies in a blood cavity.

The conspicuous grooves, which have already been
referred to as occurring on the inner surface of the
stomach, are produced by variations in height of the
epithelial cells. In this respect the epithelium agrees
with the same layer in the stomach of the lamellibranch
Pecten. |

The Digestive Gland.—This large gland has been
known as the liver, the Hepatopancreas, and the Gastric
Gland. It is now regarded as a “ pancreas,’’ but with
additional functions, such as storing pigment and fat.
The term liver should certainly be abolished and replaced
by the name “‘ digestive gland.”’

The gland is tubular, and is formed by repeated
division of the ducts which open from the stomach.
These numerous branches ramify still further, and even-
tually end blindly as caeca. The gland, lke that of
Pecten, is composed almost entirely of these caeca and
ductules, and the connective tissue which encloses each
ductule and caecum, and is therefore to be seen between
them, is reduced to a minimum. ‘There are numerous
blood lacunae penetrating between the caeca.

The appearance of stained sections is very character-
istic, for almost all the gland cells are crowded with
large, oily-looking granules, which stain vividly with
eosin (fig. 29, Dg. gr.). They are so numerous, in fact,
that details of cell structure are almost entirely obscured.
Commencing from the opening of a ductule into the
stomach, and passing towards the blind end of one of its

294 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

branches, the following changes are to be noticed in the
character of the cells. The walls are at first made up of
ciliated columnar cells (fig. 29, Dg. ¢.'), which resemble
those of the stomach. The protoplasm is distinctly seen
here, since there is little or no secreted or absorbed

substance. These cells are gradually replaced by more.

and more vacuolated ones (fig. 29, Dg. c.”), containing
the intensely staining bodies. Cilia disappear, and
finally the typical broad cells almost meet in the centre,
so that the cavity becomes very narrow, almost occluded,
as the end of a caecum is reached. —

There are usually said to be three kinds of cells
present in the digestive gland of molluscs—ferment
cells, granular cells (liver cells), and lime cells. It is
extremely difficult to make out these different types in
the whelk. Very occasionally cells are met with which
are possibly lime cells, but most cells are alike in con-
taining the oily or fatty bodies already referred to.
Whether it is really possible to draw a distinction
between ferment cells and granular cells is very doubtful.
It is probable that both are merely stages in the life of
the same cell.

Intestine.—The intestine differs hardly at all in
structure from the oesophagus. ‘There is the same layer
of very tall and narrow epithelial cells, with long cilia.
Gland cells, however, are far more numerous. The
eosinophilous cells are present in considerable numbers as
before. The lumen contains large quantities of the
granules from the latter cells, and considerable numbers
of extruded cells or parts of cells.

BUCCINUM. 295

THE ODONTOPHORE.

The complex odontophore of the whelk was examined
by Geddes in 1878. He does not describe the structures
in detail, but gives figures of some dissections.

In order to expose this organ, the proboscis should
be removed and pinned down with the dorsal surface
uppermost. If a cut is made down the middle line and
the flaps folded back, the whole apparatus lies somewhat
as figured in Pl. III, fig. 19, the oesophagus resting on
the odontophore.

Now the odontophore consists essentially of a band
(the radula), to which are attached a large number of
muscles. This band is fixed, pulley-like, on a grooved
support, which we shall call the odontophoral cartilage.
The whole structure is bound together by two delicate
sheets of transverse muscle fibres. One of these will be
seen immediately the oesophagus is removed, and lis
dorsally, forming a roof over the radula and cartilage
(fig. 19, d.m.s.). Two dorsal protractor muscles (fig. 19,
d. pr.m.) run from the anterior end of the proboscis
walls to the odontophore. In addition to these, the walls
of the buccal cavity are muscular, and there are two
delicate ventral protractor muscles (fig. 18, v. pr. m.).
It seems probable that protrusion of the odontophore (it
can be brought up to the mouth opening), is aided by
blood pressure, in addition to muscle action.

A conspicuous central dorsal muscle (figs. 19 and 16,
C.d.m.), which is attached to the extremity of the radula
(dorsally), extends back over a large number of other
muscles, all of a faint red tint, to become attached to
the floor of the proboscis. At the sides of these muscles
are two peculiar straps, consisting of a connective tissue
matrix with numerous muscle fibres (figs. 16, 18, 19, 20,

nth

296 TRANSACTIONS LIVERPOOL BIOLOGICAL, SOCIETY.

L.t.b.). These are extremely important structures, and
run back a considerable distance, to branch finally and
become attached to the floor of the proboscis. They will
be termed the Lateral odontophoral bands. These bands
form the meeting place at their anterior extremities for
a number of dorsal and ventral muscles and the odonto-
phoral cartilage. Thus many of the muscles of the
radula are not attached to the walls of the proboscis
directly, but to two lateral musculo-connective tissue
bands. Neither Geddes nor Oswald have figured this
muscle connection with the lateral bands correctly.

The odontophoral cartilage is formed of two long
band-hke lateral cartilages, which are much thicker at
their anterior ends and somewhat L-shaped in section.
They fuse ventrally at their anterior ends, and thus by
reason of their shape form the walls of a gutter or groove
open dorsally (figs. 20, 18 and 23). In addition, they
are united posteriorly by the sheet of transverse muscle
fibres, the companion structure to the dorsal sheet already
noticed (fig. 18, V.e.m.).

Upon this odontophoral caption hes the radula.
There is usually about 20 mm. of it on the dorsal surface,
and 10 mm. on the ventral.

The muscles of the odontophore may be divided

rody x0) 2——
I. Muscles attached to radula, (a) dorsally, and
(6) ventrally ;
II. Muscles attached to odontophoral cartilage ;
or

I. Protractors, (a) of Pharynx, and@(@)ia=
Radula and Cartilage.
Il. Retractors, (a) dorsal, and (6) ventral.
The protractors of the Pharynx consist of two
rauscles which run dorsally from the anterior end of the

BUCCINUM. 297

proboscis to the posterior end of the pharynx. By
contraction of these muscles the pharynx can be moved
forward.

The protractors of the Odontophore have already
been referred to. They are first a pair of muscles which
run from the walls of the proboscis to the sides of the
odontophore, really to the lateral “‘ cartilages’’ of the
odontophore, and a pair which are situated ventrally and
anteriorly (figs. 18, 21, V. pr. m.). These are also
inserted in the odontophoral cartilages. In addition to
these might be included the muscles of the buccal cavity
walls, which are attached to the radular sheath. The
action of all these muscles is to pull forward the radular
apparatus.

The Retractor muscles are much more complicated,
and are at first somewhat difficult to follow. There are
two series of these muscles, dorsal and ventral, lying
above and below the odontophoral cartilage respectively.
The dorsal retractors are much more numerous, and
larger than the ventral, and, as will be seen later, this is
to be correlated with the movements of the radula and
the arrangement of teeth on the same.

The first retractor to be observed is the most dorsal
unpaired median band (fig. 16, C.d.m.), which is attached
to the end of the radular sac, and after running back
some distance is inserted into the walls of the proboscis.

Before referring to the other dorsal retractors
attached to the radula, mention must be made of two
curious muscles which run from the end of the radular
sac, at the point of insertion of the central dorsal muscle,
to the two cartilages (fig. 16, c.c.). They are thus fixed
to two apparently unstable points. The action of these
dorsal occlusor muscles and the median dorsal muscle is
interesting, for the contraction of the former will bring

298 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the two cartilages together, closing the gutter and
preventing the radula from slipping up over the sides,
whilst the contraction of the median muscle will pull
back the radular sheath and even the pharynx.

There still remains a large number of retractor
muscles attached to the dorsal portion of the radula. Of
these, two on each side arise from the anterior ends of
the lateral odontophoral bands (or the posterior ends of
the odontophoral cartilages), and are inserted laterally
to the radular sheath (fig. 16, l.r.r.’, l.r.r."). ‘The others
all arise on the floor of the proboscis wall some distance
posteriorly. They comprise (1) the median muscle
lying under the centre dorsal muscle (fig. 16, c.d.m. inf.),
(2) the. paired muscles (fig. 16, r.’, r.”), and (3) the
paired muscles (fig. 16, V. 7.).

Altogether, there are four or five pairs of muscles
running together in this way, and all are attached
laterally and ventrally to the radular sheath some distance
forwards.

The Ventral Retractors are much inferior in strength
and number to the dorsal muscles. Like the latter, they
are attached both to the floor of the proboscis sheath
and to the anterior ends of the lateral odontophoral
bands. In fig. 18, the ventral muscles are supposed to
be seen through the odontophoral cartilage, all the dorsal
muscles and dorsal part of the radula having been
removed. It will be seen that on each side there is a
bi- or tri-partite muscle, the lateral ventral retractor,
which runs from the lateral tongue band, and is attached
in front to the sides of the radula (figs. 17 and 18, v.v.m.).
In addition to these there are two median ventral muscles
(tig. 17, m.v.r.), which lie in close contact with the
ventral sheath of circular-muscle fibres (fig. 18, v.e.m),
and then run back to be inserted in the same region as
the dorsal retractors, the floor of the proboscis.

BUCCINUM. 299

In regard to the mode of action of the apparatus,

Huxley came to the conclusion that the “‘ cartilages ’’
which support the radula were perfectly passive and
that the radula was pulled backwards and forwards,
as a strap over a pulley or a polished surface, so that it
scraped the object like a rasp or file. This was the result
of observations on some living gastropoda. Geddes held
the opposite view, a view formerly hinted at by Cuvier,
to the effect that the action of the radula was due to the
muscles pulling the whole tongue up and down.

My observations lead me to support Oswald, who
asserted that both these movements played a part; but
those described by Huxley seem, at the same time, to be
the most important.

The mere fact of the attachment of the muscles to
the odontophoral cartilage shows that this is not
altogether passive. Again, the effect of the dorsal
retractors pulling on the radula would be to cause the
cartilage to move dorsally, but this would only happen
to any extent af the radula were fixed securely against
the cartilage. Most of the muscles, however, are
inserted in such a way that much power would be lost
if they were only moving the cartilages; in fact,
it would be difficult to account for their positions.
Furthermore, there can be little play for the cartilage
in the securely bound up odontophoral mass. One would
conclude, therefore, from the anatomy alone that the
rasping movements of the radula over the cartilage were
the most important, whilst at the same time this
structure was not altogether passive. Any doubt,
however, was dispelled by one of the whelks actually
attacking the finger of the author and rasping away for
a few seconds on the skin. The median teeth of the
radula are so placed that at the point of reflection of the

300 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

latter over the tongue they point upwards. It is not easy
to see how they could be of much use in boring if they
remained in this position. One sees here perfectly clearly
the reason for the powerful dorsal musculature. When
the radula is pulled by the ventral muscles the teeth slip
backwards over the object with little friction since they
ure pointing the other way, but when the dorsal muscles
contract, the teeth are directed against the object and
much resistance has to be overcome. .

The action of the radula can, as a matter of fact, be
easily demonstrated in whelks that have been narcotised.
If the proboscis is removed from one of these animals and
squeezed between the finger and thumb gently, the
odontophore will be protruded, and the application of a
little pressure to a spot which can be found by trial will
cause the radula to move backwards and forwards over
the cartilage.

The action of the muscles, so far as I have been able
to determine, is as follows :—By contraction of the dorsal
retractors, the radula is pulled so that the teeth rasp the
object. At the same time the occlusor muscles come into
action and hold the sides of the cartilage together, so as
to maintain the groove. I find no evidence whatever to
show that by contraction of these muscles the radula
comes to lie on the sides of the cartilages as stated by
Oswald. On the other hand, the position of the lateral
teeth show the necessity of the groove, for as they pass
from the plane ventral surface of the tongue to the gutter-
like dorsal surface, they ‘‘ bite’’ inwards, and so the net
result is a tri-partite attack on the object.

The ventral muscles now come into play and draw
the radula easily backwards. There would be no point
in the arrangement of the lateral teeth whereby they can
rotate inwards and act in a most efficient manner, if

BUCCINUM. 301

movements of the radula over the cartilage were not the
usual method of attack.

The radular teeth (fig. 14) are, in accordance with
the Rachiglossan formula, three in number, one median
and two lateral. The median teeth, known as the central
or rachidian, are placed with their anterior margins
exactly transversely across the radula and possess a
number of similar denticles which will be referred to
again below. A thickened yellow band marks the
position of the tooth itself. All the teeth are fused to
the chitinous radular band. The lateral teeth are set
obliquely, and almost alternate in position with the
centrals. The inner end of a lateral tooth hes almost
opposite the base of a central tooth, whilst its outer end
is opposite the next posterior central. Furthermore, the
lateral teeth are not exactly on the same plane as the
centrals, and as the radula passes into the lingual groove
they are caused to rotate until their denticles are directed
towards the middle line. The denticles of the lateral
teeth are much larger than those of the centrals and vary
in shape, the outermost being by far the most powerful.

On the whole, the radular teeth are very regular and
characteristic in the gastropoda, and are commonly used
for purposes of classification. The teeth of Buccinum
undatum, however, make a very striking exception to
this rule, since the number of denticles on both the
central and lateral teeth varies in an extraordinary
manner, giving an excellent example of meristic varia-
tion. The three first radulas taken from Port Erin
whelks varied in number of denticles as follows :—

Lateral teeth:—4. 4. 4
Central teeth:—6. 7. 8.

Bateson records that from 27 specimens the varia-

tions were as follows :—

a ae
ee a

802 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Linteral 1- AvaiBor 4: dette (Be 4. 1316 ees
Central’ =) 52" 6.9). 6-839 9: Gs ihe 8.
Cases 258% M2 el els ah le in if i

It will be seen that even the bilateral symmetry may
be destroyed, the opposite lateral teeth having a different
number of denticles. Furthermore, though in almost
all cases the teeth are the same on the one radula, there
have been observed specimens where the number of
denticles on the anterior central teeth was less than the
number at the posterior end of the radula.

Histology of Odontophoral Cartilage
and Radular Museles.

The cartilage of the molluscan radula (fig. 23)
represents probably the earliest development of cartilage
in the animal kingdom, and by reason of its distinctness
and ease of preparation it is a good example for the study
of invertebrate cartilage.

The most external bounding layer is a delicate
connective tissue, which encloses the cartilage cells. The
cartilage itself has the appearance of a plant tissue. It
seems at first sight to be composed of very large irregular
cells with small round nuclei and extremely definite and
deeply staining cell walls. The cells (fig. 22, Cart.) are
184 in diameter, turgid with fluid, and contain a
delicate fibrillar protoplasm which does not stain
intensely owing to its attenuated state. The nuclei —
(fig. 22, Nuc.), which are perfectly spherical, are only —
4u in diameter. A nucleolus is usually present and
many small granules of chromatin. The apparent cell
walls are in reality the intercellular matrix formed by
the cells. This is small in quantity and is formed in such
a regular manner that it has the appearance of cell walls
rather than a matrix. This pseudo-wall, if followed,

BUCCINUM. 803

often appears to be continuous round two or more cells
(fig. 23). The more cells round which it is continued the
thicker it is, and hence in places there appear to be septa
running into the mass and forming at the same time the
bounding walls of contiguous cells. The cartilage of the
molluscan odontophore has often been referred to by
writers, but has apparently never been examined in any
detail. It has been compared to cellular or vesicular
connective tissue, or called cartilage without micro-
chemical tests. Josef Schaffer seems to be the only
worker who has examined it in any detail, but his paper
has appeared only as a resumé without illustrations.
The main conclusion is that the ‘‘cartilages’’ vary
towards or away from a distinctly real cartilage, and that
the development of a low or high type of cartilage is to a
certain extent independent of the phyletic position of the
animal. The Buccinum cartilage is most certainly a real
cartilage. Micro-chemically it reacts in a slight but
distinct manner to thionin, giving the characteristic
muco stain. It is therefore to a certain extent a muco-
cartilage.

The odontophoral cartilage is not entirely composed
of cartilage tissue, as the dorsal portions of the walls of
the groove (fig. 23, Ling. con.) are of a very compact con-
nective tissue, composed of a matrix resembling the inter-
celiular matrix of the cartilage area, but penetrated by
a large number of muscle fibres. |

The muscle fibres (fig. 22) of the radula are also
striking in structure. They are red in colour, and
contain much sarcoplasm. Each muscle cell is a spindle-
shaped structure of considerable length. In section it is
circular, and the diameter of these fibres is considerable,
154. The sarcostyles (fig. 22, Mus. col.), or actual
contractile elements, are arranged round the periphery of

SSR ae

304 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the muscle cell, and the large remaining central space is
occupied by the sarcoplasm. Thus in any transverse
sections of these muscles, the fibres are represented by
circular discs, the centre of which is granular proto-
plasm, whilst the periphery is either a dark staining circle
or this may have been resolved by the stain into a number
of segments—each a transverse section of a sarcostyle.
The sarcostyles do not run quite longitudinally along the
muscle fibre or cell, but wind spirally round it. Thus a
delicate double striation is present, but cannot be seen
unless one focusses completely through a fibre.

The Radula, with its teeth, is being formed con-
tinually by the cells of the radular sac in which its
posterior dorsal extremity lies, and as the teeth of the
anterior region are broken away in action, new regions
move forwards and take the place of the old. The
radular sac is a delicate cellular caecum continuous
anteriorly with the epithelium of the pharynx. It ends
blindly at the point to which is attached the centro-
dorsal retractor muscle (figs. 19 and 16, C.d.m.). The
cells forming the wall of the blind end are known as
Odontoblasts. These secrete the teeth and the basement
membrane which bears them. In Buccinum the odonto-

‘blasts are very numerous and exceedingly narrow.

Their length, however, varies in a regular manner
according to position, so that cushions of cells are formed.
Transverse and longitudinal sections are both required to
elucidate the structure of the radula sac. The transverse
section (Text-fig. 5, 4) gives the key to the structure, for
it is seen that the radula near and up to the point of
origin is so fixed that the lateral portions with the lateral
teeth are turned up at right angles to the median area.
The cells of the radula sac are longest at the extreme end,
where the radula is formed. As a matter of fact, the

‘ BUCCINUM. 305

external surface of the sac at this point forms a perfect
circle in transverse sections (Text-fig. 5).

The cells of the lateral and basal walls of this
circular sheath are of medium length. The cells of the
dorsal wall are of extraordinary size and extend down
into the cavity, forming a deep ridge, which extends for
some distance from the blind end. ‘This odontoblast
ridge lies, of course, in the gutter formed by the radula.
Thus one sees that the cells of the lateral and ventral
walls are directed towards the basal ventral plate of the
radula, whilst the dorsal ridge cells are directed towards
the teeth, both median and lateral.

Rod. ep.

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AG, With py THE | 1 |
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: aecie a ah wl 1 nt wrrracrgednneseee oe

eth Rad. ae

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As one passes towards the pharynx the depth of all
the cells decreases, the dorsal ridge passes away and
gradually the ordinary epithelial type is reached
continuous with the epithelium of the pharynx.

The nuclei of the odontoblast cells are oval, more
or less elongated and contain numerous small granules.
The most characteristic feature of the odontoblast, how-
ever, is the free end of the cell. The secretion leaves it

306 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

in the form of numerous cilia-like processes which, in a
Mallory-stained section, are at first red in colour. They
soon become blue or purple as one passes from the cell,
and then lose their individuality in a homogeneous blue
stained mass of chitin, which forms the basis of the ribbon
and teeth. The fibrous ground can be detected every-
where in young teeth stained with Mallory’s connective-
tissue stain. In the whelk the teeth are absolutely
continuous with the basal membrane of the radula, and
it is almost impossible to make out any line of junction
or to say which cells have participated only in the
fabrication of the teeth. The ventral and lateral cells of
the blind end have, however, most certainly played the
greater part in the formation of the ribbon membrane and
the dorsal cells, the teeth. |

One point, however, of great importance is that
the ventral and lateral cells are still connected with
the radula by the cila-hke tags some considerable
distance away from the blind end, and the same
applies to the dorsal cells. This probably indicates
that when the point of the radular membrane (in
this attached region) was at the extreme posterior end
of the sac, the cells below it were there too. In

‘other words, just as the radula is secreted and pushed .

forwards, so do the odontoblasts move forwards with it,
and new ones are formed at the blind end of the sac.
There are at present two views on this subject. One is
that the odontoblasts, very many of which secrete one
tooth, remain functional after that tooth has been formed
and go on secreting the next, and soon. The other view
is that the odontoblasts have performed all they are
capable of in secreting one tooth and that they pass
forwards to be replaced by new cells. Obviously they
have to become much smaller. The sections of the

BUCCINUM. 307

whelk’s radular sac favour, then, the latter theory. The
very young teeth are probably entirely formed of chitin.
Certainly there is no differentiation given by stains.
They very soon become hardened by the deposit, of
mineral salts, but they differ, as do all other Odontophora,
very considerably from the Docoglossa, where the teeth
contain a remarkably large proportion of silica. Accord-
ing to Miss Sollas, the composition of the teeth of
[nttorina is:—Ash 3°7 per cent. containing iron, calcium
and magnesium, while the rest is organic matter, the
- basis being chitin. In the Docoglossa, on the other hand,
the mineral matter may amount to as much as 27 per cent.
(Patella vulgata)—silica hydrate being present in large
quantity. |

Outside the odontoblast layer is a very compact
connective tissue layer, and it is in this connective tissue
that the muscles, which are attached to the radula,
terminate. The muscles are of the type described above.
Their terminations can be followed very beautifully
indeed in sections stained in Mallory, for the muscles are
bright red, and the connective tissue bright blue.

BLOOD VASCULAR SYSTEM.

The vascular system of some molluscs has attained
a high degree of complexity. This is especially the case
with the Cephalopoda and some of the Prosobranchiata,
and the whelk amongst the latter may be taken as showing
a good example of a well developed molluscan blood
system. It seems that, at the present day, in many
zoology courses there is a tendency to pay little
attention to the vascular system of the molluscs.
The cephalopods receive perhaps adequate treatment,
and possibly the snail (Helix) has some attention.

308 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

In most cases, however, beyond the heart and
large vessels leaving it, very little is investigated. Now
in the whelk it is quite easy to demonstrate all the more
important facts in connection with the circulation, and if
a little care is used in injecting, there is no reason why
this method should not be adopted in the ordinary
laboratory classes for senior students. Very few complete
accounts have been given of prosobranch blood systems,
and it is hoped that this description will serve as an aid
in the study of an excellent and exceedingly common
type.

A great part of both the venous and arterial systems
can be made out from the surface, without any dissection.
All the vessels shown in figure 35 on Plate V can be seen,
with the exception of the dotted trunks which run in the
foot. This means that a cold injection mass may be used,
without troubling to employ any of the complicated and
unsuccessful mixtures which have been invented in order
to attain solidification or coagulation in the vessels.

Directions for Injecuvom

For students’ purposes, two very simple injecting
masses will suffice :— mi

1.—Cold Injecting Mass.—Use one of the familiar
collapsible tubes of artists’ oil colours. Chrome-yellow 1s
a convenient tint. Squeeze this into a dish and dilute with
turpentine, stirring until a slightly thick but uniform
mixture is produced.

2.-Hot Injecting Mass, for studying the deeper
vessels.—Melt down one or two pieces of ordinary table
jelly in an evaporating dish, adding water 1f necessary.
Add to this some carmine, rubbed down with a small
quantity of water in a mortar and stir until a uniform
mixture is obtained which will solidify on cooling.

BUCCINUM. 309

It is absolutely essential that the whelks to be
injected be dead, and further that they have died in a
thoroughly lax uncontracted state. The best means is to
allow the whelks to expand in a small quantity of sea-water
and then add carefully a few drops of a 2 per cent. solution
of cocaine in 90 per cent. spirit; and continue adding
gradually a little of the cocaine solution. It will be
found that three days are necessary before the animals
become irresponsive to stimuli. By that time the water is
probably in very bad condition, but the whelks will not
be seriously affected by this, so far as injection is con-
cerned. The shell should now be removed piece by piece
with a pair of bone forceps, and great care must be taken
that the soft tissues are not damaged. The columellar
muscle should be detached from the shell with a scalpel.

On the external surface the position of the mucous
gland and the gill should be made out. Those two organs
of the pallial complex with the osphradium can be seen
through the mantle. The pericardium has been already
seen, lying at the side, and under, the kidney.

Injections can be made from three places.

1.—The syringe should be inserted into the efferent
branchial artery, with the point towards the heart. This
vessel can be seen quite easily, forming the ventral
boundary of the gill. If the paint mixture is used, a very
clear view will be obtained of the branchial vessels, the
vessels of the mucous gland, and the reno-mucous vessel
(fig. 35, Mu. gl., Br. v., R. sim.). The auricle will be
filled, and probably the efferent renal vessels (fig. 35,
Ren. eff.).

2.—The point of the syringe should be plunged inte
the foot, so that it reaches somewhere in the space in
which the alimentary canal and poison gland lie. This
injection will fill the extensive venous sinus, and then

310 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the veins bringing blood back from all parts to the renal
organ. They can be seen by removing the auricle and
ventricle from the pericardium, and by slitting the renal
organ along its left ventral margin and thus laying bare
its internal surface.

3.—The syringe should be inserted into the aorta
(figs. 35, 36, Ao.) at the point where it leaves the ventricle.
This injection should be performed with the hot jelly, but
it is the most difficult one and often fails. The arterial
system is injected by this means.

THE HEART.

Buccinum possesses (as do all Monotocardia so far as
is known, except Cypraea) only one auricle (the left one
of the lower Gastropods).

The auricle and ventricle lie in the pericardial cavity,
which is bounded by the renal organ and the digestive
gland, the auricle being anterior.

The auricle is a very thin-walled sac, capable of
considerable extension. It 1s somewhat like a pear in
shape with the pointed end situated ventrally, and into
this end opens the efferent branchial vessel.

There are two other openings into the auricle which

~ are situated at the dorsal end. One of these at the anterior

edge of the pericardium is the opening of the nephridio-
cardiac vein, the other, which is situated on the posterior
surface, is the opening into the ventricle. The latter,
the auriculo-ventricular passage, 1s guarded by a valve
so that blood is prevented from passing backwards from —
ventricle to auricle. 3

Tt will be noticed that the nephridio-cardiac vein
opens directly into the auricle and not into the efferent
branchial vessel. It is often somewhat difficult, however,
to determine whether the vein enters near the opening of

BUCCINUM. alk

the branchial vessel and lies along the anterior margin,
or whether it enters more dorsally.

The ventricle is very different in appearance from the
auricle. It is roughly spherical, with very thick spongy
walls, but the cavity is so much reduced by crossing
muscles that the consistency of the whole is very like that
of a sponge. Injections into the ventricle hardly ever
succeed, because most of the injecting fluid oozes out at
the point of insertion of the syringe.

ARTERIAL SYSTEM.

From the ventral pole of the ventricle a single vessel
arises, the Aorta (figs. 35, 36, Ao.). This is of very short
length for it divides almost immediately into two branches,
the anterior or Cephalic aorta (figs. 35, 36, A.c.), and the
posterior or Visceral Aorta (figs. 35, 36, A. vis.). The
Anterior Aorta gives off immediately a small vessel
(fig. 36, a.’) which sends branches to the oesophagus and
the columellar part of the spire and then plunges below
the floor of the mantle cavity into the large sinus in which
he the oesophagus, salivary glands and proboscis. The
aorta lies at first at the left side but soon crosses over the
oesophagus and runs under the salivary glands. Just
after entering the body cavity, it gives off a vessel on the
right side which passes to the columellar muscle (fig. 36,
A. col.) and branches on its surface. The next large
vessel leaves the under surface of the Cephalic Aorta
(fig. 36, A. pall.) and passes through the muscles to
reach the surface of the mantle. It divides into two main
trunks, of which the ventral one reaches the surface on
the under side of the animal, and the dorsal vessel just
below the osphradium (fig. 35, A. pall.’).

This dorsal pallial artery supplies the siphon, and

a |

312 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

runs round the edge of the mantle, giving off branches on
both sides. Very small vessels leave the aorta at intervals
and pass to the alimentary canal. The next two vessels —
which arise are asymmetrically placed. They leave the
aorta laterally, but the left one 1s somewhat posteriorly
situated. These vessels (fig. 36, A. cut.), after passing
out laterally, run up the walls of the body cavity and
break up into small vessels innervating the roof (the floor
of the mantle cavity). The vessel on the right side,
however, gives off an additional branch (fig. 36, A. cut.’),
which bores into the wall and passes to the tissues below
the ovary and rectum.

The Cephalic Aorta passes forwards until the Nerve
Collar is reached and then breaks up at one point into
several vessels. The point where this division takes place
is hidden by the salivary glands and by connective tissue.

One large vessel runs down at right angles to the
course of the Cephalic Artery. This is the Pedal Artery
(figs. 86 and 35, A. ped.). After a short course, it divides
into two branches which make their way into the foot,
branch again (each in a similar way) and supply the
musculature. These blood vessels are of considerable size.
Here they branch, but most of their small branches, which

form almost a capillary network, are not shown in the

figures. It is by means of this network, and the forcible
action of the blood, that the great expansion of the foot
is effected.

Returning to the nerve collar region, it will be seen
that another large vessel runs dorsally, also at right
angles to the aorta. This artery, the Buccal (fig. 36,
A. buc.), goes forward at the side of the oesophagus, and
after giving off two small vessels to the proboscis, enters
the latter with the alimentary canal. Here it breaks up
into a complicated series of branches supplying the

BUCCINUM. ols

odontophore and the muscles. Two other prominent
vessels (fig. 36) arise at the nerve collar, one on either
side of the aorta. They both pass toward the tentacles but
before reaching them, small vessels leave dorsally to
supply the tissues of the “head”’ (fig. 36).

The right tentacular artery gives off at its origin a
- large branch in the male, the penis artery (fig. 36,
A. pen.).

The Visceral Aorta (figs. 85, 36, A. vis.) can be seen
quite distinctly without injecting. It turns abruptly
after leaving the ventricle and runs close to the surface
across the intestine. Half-way across it gives off a branch
(figs. 35, 36, A. g.) on the right side, which passes to the
stomach and breaks up into numerous twigs upon its
external surface. The main trunk plunges into the mass
of the digestive gland, immediately after crossing the
intestine, and runs right through, to appear again at the
surface on the other side. It now remains at the surface,
just below the epithelium, and lying upon the digestive
gland a little ventral to the edge of the gonad; and in
this position runs to the end of the visceral mass.
Branches are given off at intervals which run out at right
angles to supply both the gonad and the digestive gland
(fig. 36, A. go.).

Tuer Venous SYSTEM.

The blood which is carried to all parts of the body
by the arteries, collects in lacunae and is brought back to
certain more extensive central sinuses. Many of the
paths which are taken are, however, so narrow and so
well-marked that there almost appears to be a definite
system of capillaries connecting up the arteries and veins.
These channels are mere excavations, which may be lined

3814 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

by a pseudo-epithelium due to modified cells. The centre
of the venous sinus system is the region on the surface of
the digestive gland at the back of the pericardium and
underneath the renal organ (fig. 37). There is a large

sinus—the largest in the body—underneath the pallial
cavity (fig. 37), while an important vein runs along the
right margin of the mucous gland (figs. 35 and 88,
R. Sin.). These important regions may be named as
follows:—The sinus in the anterior perivisceral cavity
will be termed the Cephalic Sinus (fig. 37, Ceph. Sin.),
as it collects blood from the head and foot. The branched
system, situated at the back of the pericardium, and
under the renal organ is the Kenal Sinus System (fig. 37),
while the long vessel-like sinus which extends from the
right side of the renal organ to the anterior end of the
mucous gland is the Reno-Mucous vessel (figs. 38 and 30,
Fes ‘sity.

Blood returning from the viscera (digestive gland
and gonad) passes by means of superficial vessels (fig. 35)
to a sinus which extends along the columellar surface of
the visceral mass. This sinus stretches as far as the
region of the pericardium and there becomes connected
with the renal sinus system (fig. 37), by means of which

‘the blood reaches the renal organ.

Blood returning from the head, proboscis, and foot
flows into the great cephalic sinus (fig. 37, Ceph. Sin.),
and from here two paths may be taken. These leave the
perivisceral cavity at the posterior extremity of the
mantle cavity, and are close together. One ascends and
reaches the renal organ directly. It passes along its left
side, internally, so that it cannot be seen unless the renal
organ is turned back as in figure 37. The other receives
blood from the oviduct and rectum (figs. 37, 38) and enters
the Renal Sinus System and so to the renal organ.

BUCCINUM. 315

THe Renat Buoop VESSELS.

It will be convenient now to discuss separately the
vessels in the renal organ. This organ forms a junction
system of small vessels which is interpolated into the
venous system. The blood which enters it may leave by
two paths, one of which goes to the heart directly and the
other by way of the gills.

The blood vessels of the renal organ of Buccinum are
remarkably organised. This, however, is probably
general in the higher prosobranchiates. or purposes of
comparison it will be advisable to use the renal organ of
the Lamellibranch Pecten, which has been caretully
worked out. In this genus the renal organ is a tube, the

inner wall of which is thrown into folds in order to ~

increase the area of renal epithelium. Between these
folds are blood cavities. The blood leaves the renal organ
by one path only and then passes to the gills.

In Buccinum, the renal organ is a similar tube, but it
is erescentic in section and the inner wall of the outer
half only is thrown into folds. The folds are quite
different from those in Pecten (see Renal Organ below)
and the floor of the renal organ (the unfolded side) covers
a large vessel of the Renal Sinus System (fig. 37). The
folds of renal epithelium hang down into the lumen of
the renal organ, and from them membranous extensions
pass to the floor. Now, on the floor, we have the large
vessel of the Renal Sinus System, and from it vessels
arise which pass through the lumen of the renal organ to
the folds of renal epithelium.

Thus a large quantity of blood enters the renal organ
over the internal surface. This is a most curious position
which a glance at the diagrammatic section (fig. 51, 4.)
will make clear.

316 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

This, however, is not the only path for blood
entering the renal organ. We have already seen that a
large vessel runs round the margin (fig. 37, lam. v.).
From this vessel branches arise which run across in the
plane of the outer surface and divide up close to this
surface. The system of anastomosing channels is
extremely complicated, and minute, and finally all
is resolved into two outgoing paths. One of these
is seen right on the surface when the renal organ
is injected, and is shown in fig. 35. It consists of
delicate vessels running almost at right angles to
the left margin (fig. 35). These delicate vessels
enter a large lymph-like gland, the nephridial gland,
which extends along the side of the renal organ against
the pericardium. It was supposed that the nephridial
gland communicated only with the auricle and that blood
passed to it by means of the nephridio-cardiac vein and
back again by the same course. A detailed study of this
point in the whelk has clearly shown that there is a direct
path from the renal organ through the nephridial gland
to the heart. The other path is more internal and is
made up of another series of small vessels which open into
the reno mucous vessel running along the right side of the
renal organ (fig. 38). Thus there are two lines of com-
munication bringing blood to the renal organ and two
paths along which blood leaves.

Blood. Cireulation in Pallial @ompiee

The blood from the ‘‘ kidney ’’’ which pours into the
reno-mucous vessel passes forwards into that part of the
latter bounding the mucous gland. Blood from the rectal
region and mantle also enters this vessel, and from it
passes through the mucous gland to the afferent
branchial vessel. The mucous gland is exceptionally

BUCCINUM. SEC

well suppled with blood, as will be seen from figure 35.
From the afferent branchial vessel a large number of
filamental vessels arise which convey the blood through
the gills to the efferent branchial vessel. The efferent
branchial vessel collects some blood from the mantle
which has passed through neither gills nor renal organ,
but the quantity is small. Finally it enters the auricle.

To summarise :—

1. Blood passes from the ventricle to various parts
of the system.

2. Blood is collected into sinuses and conducted to
(a) Renal Organ, and (6) Gills.

3. The blood passes back to auricle through (a)
Renal organ and gills, (b) Renal organ alone, (c) Gills
alone, and (d) Mantle alone.

The largest quantity of blood passes through routes
(a) and (b), and that passing by method (d) is almost
negligible.

THE NERVOUS SYSTEM.

Once more we must emphasise the fact that we are
dealing with an example of the most highly developed
Prosobranchs. The Nervous System presents the twisted
visceral loop characteristic of the Streptoneura, together
with extreme concentration of the ganglia. So far
as the description of the nervous system is concerned,
constant reference has been made to the excellent work of
Bouvier on the nervous system of the prosobranch
gastropods. All the important details have been worked
through again, however, and the figures differ in some
minor details from those given in Bouvier’s memoir.

Owing to the extreme concentration which has taken
place, the nerve ganglia may be divided into two groups.

318 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The anterior centres are situated in the cephalic region
and surround the alimentary canal, forming a collar
almost hidden by the salivary glands. The posterior
centres lie just behind the pallial cavity in the region
between that and the visceral mass. The two groups of
vanglia are connected by the long visceral com-
missure (fig. 41, vis. com.). Dissection of the nervous
system is a matter of some difficulty and is usually a
stumbling block to some students. It is quite easy to
find the nerve collar, but rather difficult to expose clearly
the component parts owing to the presence of a tenacious
fluffy connective tissue which obscures all.

The nerves and commissures passing into the body
wall are also difficult to follow by reason of the toughness
of their surroundings and their resemblance in colour to
the tissues in which they are embedded.

It is best to dissect the nervous system in whelks that
have been kept for a short time in alcohol. This hardens
the muscle but brings out the nerves much more clearly.
In addition, the action of a solution of oxalic acid is
decidedly useful. It brings out the otocysts and clears up
the muscular mass of the foot. In order to investigate
the centres making up the nerve collar, it is best to cut

through the cerebral commissure which lies above the

oesophagus and then to dissect the alimentary canal care-
fully away. The ring of ganglia can be examined from
the wmside, and thus observed without requiring the
removal of so much connective tissue.

The Anterior Centres are ten in number. They consist
of the cerebral, pleural, pedal and buccal ganglia, which
are paired and the supra- and sub-intestinal gangla
which have been drawn in, in the general concentration,
until they have reached a position close to the pleural
ganglia (fig. 42, Sup. ut. and Sub. int.).

BUCCINUM. 319

The Pedal Ganglia are the largest ganglia in the
whelk and are situated most anteriorly. They are oval
in shape and lie upon the floor of the perivisceral cavity,
in close contact. Consequently there is no pedal com-
missure (fig. 42, ped. g.). In the female these ganglia are
symmetrical, but owing to the origin of the large penis
nerve from the right ganglion, in the male, they become
asymmetrical (fig. 43). The pedal ganglia are connected
to the cerebral and pleural ganglia on each side by two
very short connectives (fig. 42, ¢.p., pl. p.), a small space,
the “‘ triangle lateral,’’ being left between the ganglia
and their connectives (fig. 42, trz. lat.).

A large number of nerves arise from the anterior
end of each pedal ganglion and run forwards together for
a short distance to plunge into the muscular foot. These
are the pedal nerves (figs. 41, 42, ped. n.) and they
innervate the entire foot. Other smaller nerves leave the
pedal ganglia on both the dorsal and ventral surfaces.
According to Bouvier there are two such dorsal (fig. 42,
do.) and three ventral nerves (fig. 42, ve.) from each
ganglion. The former pass to the head region whilst the
latter innervate both the foot in the immediate vicinity,
and the floor of the anterior body cavity.

The penis nerves leave the right pedal ganglion at the
posterior lateral corner (fig. 43, P.) and consist of two
trunks, one of which is extremely large and follows the
vas deferens inside the penis to form a network from
which delicate branches innervate the muscles of the
organ.

The Cerebral Canglia will be seen somewhat out of
their normal position if the dissection is conducted as
described above, for after the severing of the commissure
and the removal of the alimentary canal, they will be
turned back as indicated in fig. 42, c.

320 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

They are somewhat irregular in shape and are
connected with each other by a commissure which passes
over the alimentary canal. Owing, however, to the
folding of the gut and the almost vertical position of the
oesophagus at this point, the cerebral commissure lies
rather posteriorly to the gut. Both the oesophagus and
branches of the anterior aorta are encircled by the collar
formed by the cerebral ganglia, cerebral commissure, and
pedal ganglia. |

It will be noticed that the cerebral ganglia in the
whelk are by no means the most anteriorly placed. They
he some little distance behind the pedal ganglia and
almost directly above the pleural centres. This position,
which may give rise to a little confusion at first, is due to
the course taken by the oesophagus between the proboscis
and the gland of Leiblein. The cerebral centres are
joined by very short connectives to the pedal and pleural
ganglia, and at this point it may be observed that there is
a slight asymmetry in the relative position of the right
and left ganglia. Two other small ganglia, the buccal
(fig. 42, bucc.) are in close contact with the cerebral and
are themselves connected by a long and delicate
commissure passing in front of the alimentary canal.

Three nerves innervating the sense organs leave the
external face of each cerebral ganglion. The largest of
these is the tentacular nerve (fig. 42, tent.). It can be
followed quite easily in the tentacles, and bends at right
angles when above the ganglia to reach them. A few
twigs are sent to the head region from this nerve before
it finally penetrates the tentacle. The other two nerves
pass directly, the one to the eye, and the other to the
otocyst (fig. 42, op. n., and n. ot.). The two otocysts are
situated immediately below the anterior ends of the pedal
ganglia and just imbedded in the muscular floor. They

BUCCINUM. 321

are very easily overlooked, even when one examines the
exact place where they occur. They are about two milli-
metres apart and 0°5 mm. in diameter.

In addition to these sensory nerves, there are two
other groups which have their origin in the cerebral
ganglia, i.e., the nerves of the proboscis and of the
cephalic integument. The former arise just anterior to
the cerebral commissure and form a compact bundle on
each side, ascending with the salivary ducts and running
along with them towards the proboscis. They innervate
that organ and also the proboscis sheath. The cerebral
nerves of the proboscis enter into close relation (forming
anastomoses) with certain nerves from the small buccal

ganglia, and all pass together into the proboscis. The
buccal nerves innervate the oesophagus and radular
apparatus, the cerebral nerves on the other hand appear
entirely concerned with the walls of the proboscis itself.
There are two other nerves innervating the integument of
the head region and these arise from the cerebro-pedal
connectives (Text-fig. 6, ceph. n.). They eventually
branch, sending numerous twigs to the region immediately
posterior to the tentacles.

The Pleural Ganglia are asymmetrical owing to the
positions taken by the supra- and sub-intestinal ganglia

322 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

(fig. 42, sup. emt. and sub. int.) in the general concen-
tration of nerve centres. They are attached to the
cerebral and pedal ganglia by short connectives, the
cerebro-pleural (fig. 42, c. pl.) and the pleuro-pedal
respectively (fig. 42, pl. p.). The pleural gangla form
part of another nerve collar owing to the position of the
sub-intestinal ganglion. This, normally attached to the
left pleural ganglion, is here in addition fused to the
right pleural so that it comes to lie between the two.

The left pleural ganglion gives origin to the siphon
nerves (fig. 42, Siph. n.), the columellar nerve, and
several others which innervate the body wall (fig. 42).
The two siphon nerves pass outwards along the floor of
the perivisceral cavity until the body wall is reached.
They plunge through this tough muscular wall and arrive
at the siphonal region of the mantle. A number of
branches are given off to the siphon, and certain fibres
form important connections with the pallial nerve from
the supra-intestinal ganglion, eventually taking part in
an anastomosis in the mantle (fig. 41). A left zygoneury
is thus formed (fig. 41, zyg.).

The columellar nerve leaves the ganglion posteriorly
and crosses over the floor of the anterior cavity towards
the right side. It is not a very large nerve and is easily
overlooked. Eventually it reaches the columellar
muscle. |

The remaining nerves are small with the exception
of one which arises on the pleuro-pedal connective,
divides soon, and innervates the walls of the body cavity.

The right pleural ganglion has only one nerve of
importance here, and this corresponds to the last nerve
mentioned above, arising from the left pleural ganglion.
It takes its origin on the pleuro-pedal connective (Text-
fig. 6, r. pl. n.) and innervates the right lateral walls of

BUCCINUM. 323

the body cavity. The branches of this nerve are quite ©
easily seen if the nerve collar in situ is pressed slightly
over to the left side. The branches pass out directly to
the right.

The Supra- and Sub-intestinal Canglia. Three
prominent nerves are easily seen crossing the floor of the
anterior cavity and entering the wall on the right side.
The most anterior of these is more delicate than the other
two. All arise in the sub-intestinal ganglion (fig. 42,
sub. wnt.). The two larger are the pallial nerves (figs.
41 and 42, pall. n.), and after plunging through the walls
of the body cavity they reach and divide up on the
mantle, the more anterior of the two forming an
extensive network.

The branches of these pallial nerves of the right side
come into contact with the branches of the left palhal
nerves. This takes place in the mantle immediately
above the palhal cavity. Thus a continuous network is
formed running around the mantle edge.

A number of other small nerves (fig. 42) arise in the
sub-intestinal ganglion and innervate the walls of the
body cavity and the columellar muscle.

The largest nerve leaving the sub-intestinal ganglion
arises from the left posterior corner. It is the Visceral
Commissure itself (figs. 41 and 42, Vis. com.). This
cord runs posteriorly almost in the middle line, in close
contact with the floor of the body cavity, for some
distance, and then passes underneath the most superficial
muscle fibres. It is often very difficult to dissect out the
visceral loop in the whelk, but if the specimens have been
preserved in spirit, the track of the visceral commissure
is visible as a delicate ridge on the floor of the body
cavity running back from the point « in the figure.

The visceral ganglia are situated on the loop just

394. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

posterior to the mantle cavity. They can be seen from
the exterior, without any dissection, under the epithelium
of the area between mantle, digestive gland and
columellar muscle. The other part of the visceral loop
connecting the visceral ganglia with the supra-intestinal
ganglion is by no means easy to follow, for it lies under
the superficial muscles of the left wall of the body cavity.
It leaves the wall near the point of entrance of the two
left pallial nerves (figs. 41 and 42, pall n.”), and runs —
along with them to the supra-intestinal ganglion. The
three nerves have thus to cross over the alimentary canal
and gland of Leiblein. Of the usual figure of eight
formed by the visceral commissure, only the lower,
posterior, loop is of any extent.

The two large left pallial nerves arise in the supra-
intestinal ganglion, and passing over to the left side,
plunge through the body wall. They are extremely
close together at first, and form one thick band. On
reaching the mantle they proceed to divide. The most
anterior nerve innervates the osphradium. In addition
to this, it gives rise to some twigs which, by means of
their anastomosis with the siphonal nerve, set up the
zygoneural connection already mentioned as existing on
the left side. Both the anterior and posterior pallial
nerves from the supra-intestinal ganglion take part in
the innervation of the gills.

An important nerve arises from the sub-intestinal
part of the commissure itself, just at the posterior end of
the perivisceral cavity. It runs outwards, underneath
the vas deferens in the male (fig. 41, com. n.) to the
rectum and mucous gland.

The Visceral Ganglia are two in number. Of these
the right is much larger than the left, and gives origin
to most of the nerves. The principal nerves are the

BUCCINUM. 325

following:—The Visceral nerve (fig. 41, wvisc.) passes
close to the left side of the vas deferens in the male and
innervates the digestive gland and gonad. It is an
extremely long nerve, in fact, the longest in the animal,
and can be traced to the tip of the visceral mass. Two
nerves, the recto-genital and rectal, (fig. 41, rec.’ and
rec.) pass out to the right, to the rectum and gonoducts.
A larger nerve, the reno-cardiac (fig. 41, 7.c.) passes out,
breaking up very considerably on its way and sending a
branch to the pericardium and heart. From the smaller
ganglion a nerve arises which passes to the efferent

branchial vessel (fig. 41, eff.).

THE SENSE ORGANS.

The sense organs may be divided into simple and
compound, the former class including only the numerous
sense cells which occur scattered amongst the ordinary
epithelial cells, the latter the Eyes, Osphradium, and
Otocysts.

THE HYE.

The eyes are two in number and are situated at the
base of the tentacles, on the dorsal surface of a small
lateral protuberance.

They are visible as two round intense black spots,
but when the tentacle is contracted and the body wall is
thrown into folds, the eye, hidden in the angle between
tentacles and head, is not easily seen. No metallic
glitter so characteristic of the Pecten eye ever occurs,
and, as will be seen below, the layer in the eye responsible
for this feature is not present.

The lowest gastropoda possess eyes which are simply
sac-like invaginations of the outer epithelium. At the
bottom of the sac the epithelium is modified somewhat

326 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and forms the retina. The cavity remains open to the
exterior.

In the higher gastropods the open pit is succeeded
by a closed vesicle, and this ontogenetic sequence is

probably an indication of the phylogeny. As we should

expect therefore, the Buccinum eye is a closed vesicle cut

off from the surface. The general epithelium extends _

over it, forming an outer cornea (fig. 47, Cor.). The wall
of the vesicle is formed of a single layer of cells, once
part of the bounding epithelium of the head. These
cells are modified in various regions. The area immed-
iately below the outer cornea, through which the light
has to pass, is formed of cells free from pigment—the
area itself is the inner cornea (fig. 47, Ps. cor.). The rest
of the vesicle wall forms the retina and is made up of
pigment-containing cells.

The cells of the outer cornea differ only from
those of the epidermis in shape. As the optic
vesicle is approached, the deep and narrow epidermal
cells with slender nuclei become less deep and of
greater thickness. There is a well-developed cuticular
margin present. With this change there is an alteration
in the form of the nucleus, a spherical shape taking the
place of the slender compressed form. ‘The optic vesicle
is imbedded in a mass of connective tissue (fig. 47,
Con.) and muscle fibres, in which numerous blood
spaces occur irregularly. This tissue, which is usually
more compact near the surface, is continued in so as to
form a layer between the vesicle and the outer cornea.

The cells of the optic vesicle appear to be of
two kinds, and only two kinds of cells make up the
entire vesicle—retinal and corneal areas inclusive. In
sections, the retina, or area directly opposite the cornea,
is built up of very large cells, which can be distinctly

all

Ss —-S—ti( PC

-
;
,

BUCCINUM. 397

seen extending from the connective tissue to the cavity
of the eye (fig. 47. Met. c.). They possess spherical
nuclei in which may be seen a nucleolus and numerous
granules, and their distal portions are lost in a mass of
dense black pigment.

Between these large cells are to be found a number of
compressed nuclei (fig. 47, Nuc. int. C.), which stain more
darkly than those above mentioned, possess finer granules,
and in short cannot be mistaken for the nuclei of the
large cells. These nuclei belong to slender fusiform cells
which are interpolated between the larger ones and whose
boundaries are difficult to follow in sections. If the eyes
are macerated in a 4 per cent. cocaine solution in sea-
water, the true shape of the structures building up the
retina becomes at once apparent. The large cells possess
the extraordinary shapes indicated in fig. 48. It will be
noticed that they are widest towards the base, and that
the nucleus is in general situated not far from that region,
in fact, somewhere near the centre of the cell. One of
the most curious facts is the prolongation of the cell into
processes, of which there may be several (fig. 48, pr.).
The apex of the cell is rounded and contains a large
quantity of black pigment (fig. 48, Prg.) in the form of
minute granules closely crowded together. Very often,
too, this rounded distal extremity bears a prominence
which may be taken as the remains of the ‘‘ Stiftchen ”’
or hair-like processes seen in sections and mentioned
below. The narrow cells also possess pigment, so that
there is no differentiation here into pigmented ond
pigmentless cells.

The cavity of the optic vesicle is filled with a
structureless gelatinous mass, which forms a lens (fig. 47,
Lens). This lens does not possess a perfectly rounded
contour where it abuts on the retinal cells, but is indented

898 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and in each indentation there is a delicate bundle of hair-
hike *‘ Stiftchen ’’ (fig. 47, St. s.), which appear to break
away from the cells in macerations. Thus the eye of
Buccinum agrees with the long series of eyes possessing
Stiftchen-bearing cells. The presence of such processes
is one of the most striking characters-in the structure of
visual organs.

There has been no small difference of opinion as
to which are the visual cells in gastropod eyes. In
many cases the difference between the cells which has
struck the observer has been the presence or absence of
pigment. Here both sets possess pigment. Could
the actual nerve connections be demonstrated, the
matter would be easily solved, but unfortunately the
fibrous processes of both cells merge into the tangled mass
of nerve fibres and connective tissue below the eye.
Judging from structures in the eye of Pecten, and
especially from the character of the nucleus, the author
would say that the large cells are the visual cells not-
withstanding the fact that the cell body of the slender
cells is more in accordance with expectations.

This view is, however, supported further by two
facts. The. large cells appear to possess the ‘* Stiftchen.”’
Both kinds of cells occur laterally almost up to the inner
cornea. There the large cells begin to disappear, whilst
the nuclei of the others are very like, one might almost
say the same as, those found in the cells of the inner
cornea itself. According to this view, therefore, the
slender cells are merely supporting cells, and they are
continued across the eye to form the inner cornea.

The nuclei of the inner corneal cells are flat
compressed structures (fig. 47, Ps. cor.), like the nuclei
in the retina, but instead of being arranged at right
angles to the wall of the cavity they have rotated, so

BUCCINUM. 329

that they now lie end to end and parallel with the plane
of the cornea. The gelatinous lens is probably secreted
by the supporting cells. The optic nerve breaks up
underneath the optic vesicle, and a network of fibres
extends below the retina.

So far as observations go, the whelk relies but little
on its visual organs. It is difficult to understand how
complicated organs of this kind could be produced in the
molluseca, when one thinks of the feeble responses to
experiments. Hither our experimental methods have
failed so far to demonstrate the utility of many of the
molluscan eyes, or some of our conceptions of evolu-
tionary processes require modification. The tendency to
take anthropomorphic views, however, in work both
structural and experimental, for invertebrate sense
organs is perhaps the danger to be avoided.

THE OSPHRADIUM.

The sense organ known as the Osphradium, situated
usually in close proximity to the respiratory organs,
attains a degree of complexity in Buccinum which is
probably never exceeded in the Mollusca. In the
Lamellibranchs the osphradium is merely an area of
somewhat thicker epithelium, whose presence is not even
marked by pigment (except in Arca). It is, therefore,
quite invisible to the naked eye.

The lower gastropods, the Diotocardia, possess an
osphradium which seems to be somewhat of the same type
so far as general structure is concerned. As one passes,
however, towards the more highly developed Monoto-
cardia, the osphradium takes the form of an axis bearing
on each side a large number of leaflets. It becomes, in
fact, bi-pectinate, and resembles superficially a gill.

330 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The osphradium of Buccinum is usually obvious as
soon as the shell is removed, for it can be seen through
the mantle. There is no difficulty in finding it when the
mantle cavity is opened and the roof examined. It
stands out as a deep brown-black organ, situated right
across the gutter of the pallial siphon and between this
organ and the gill (fig. 8, Osph.). It lies, therefore, on
the opposite side of the ctenidial axis, and by réason of
its build was formerly known as the “ false gill.” It is
long, and is made up of about 90-100 leaflets arranged
on both sides of an axis, which is nothing but an
elongated nerve ganglion covered by epithelium. These
leaflets are largest in the centre and diminish in size
towards the ends, giving the whole the appearance of an
elongated oval. The leaflets are roughly triangular in
shape, those of the right or ctenidial side being, however,
larger than those of the left (fig. 44). Hach leaflet has
an inferior free edge (fig. 44, /nf.), a lateral edge (fig. 44,
Lat.), and what must be termed the third side of the
triangle—the internal edge—which is curved and fused
to both mantle and nerve axis (fig. 44, Pall and JN. az.).

The entire surface of the lateral faces is pigmented,
but the pigment is often free from the inferior edges of
_ the leaflets, a condition the reverse of that described by
Bernard for Cassidaria. For a further study of the organ :
reference must be made to sections.

Each Osphradial leaflet is a fold of the epithelial
layer bounding the inner surface of the mantle,
together with certain connective supporting tissues.
Just as in the branchial leaflets, there is an external
bounding epithelium resting upon supporting mem-
branes and leaving a series of flat blood spaces,
which are here occupied largely by branches of the
osphradial nerves (fig. 45). The epithelium resembles

BUCCINUM. oor

that of the ctenidial leaflets further in being modified
and specialised in different regions. These regions are
(a) sensory, (b) glandular, (c) ciliated, the former being
by far the largest in extent. The sensory region extends
over the greater part of the free lateral surface of the
leaflets. The glandular region is confined more or less
to the edges, and in particular to the lateral edges
(fig. 44, Osp. gl.). The ciliated area is a long narrow
strip extending along both sides of the leaflet against the
glandular edge and between it and the sensory area
(fig. 44, Osp. cil.). Thus in a section taken through
A-B (fig. 44), the glandular area is nearest B, that is
where the free lateral edge meets the mantle, this is
followed by the ciliated region, and then the remainder
is sensory. ‘The sensory area is characterised as in the
lamellibranchiata by an absence of cilia, and a con-
siderable thickening of the epithelium. This thickening
is due to an increase in the length of the cells, which are
further modified by the possession of a yellow granular
pigment. Now the real structure of this important area
is not easily investigated. Sections well preserved and
fixed show that it is built up of a large number of
pigment-bearing cells, whose nuclei are apparently
arranged at different levels because one sees nuclei close
to the basal membrane, and from here to near the free
surface they are irregularly scattered (fig. 45, Osp. so.).
There is, however, a stratum close to the free margin of
the cells perfectly free from nuclei, and it is here that the
pigment is situated.

Macerations of the osphradium explain the structure
instantly. The pigment-bearing cells are of the type
indicated in fig. 46, and all agree in possessing a terminal
plateau in which the pigment lies. From this plateau
the cell becomes constricted and fibre-like, with, however,

332 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

one large swelling where the nucleus lies. The position
of the nucleus varies considerably, and so it is possible to
pack together mary more cells than would be the case if
the nuclei were all situated at the same level.

These cells are the indifferent cells of Bernard. One
feature that he has overlooked is the well-defined cuticular
margin (in sections), crossed by delicate striae arranged
perpendicularly to the free surface of the epithelium.

Now, in addition to these cells, there are, according
to Bernard, neuro-epithelial cells which are in connection
with the nerve fibres. These neuro-epithelial cells are
always fusiform and possess a rounded or oval nucleus.
If the osphradium is an important sense organ (and one
would assume so from the size of the nerves innervating
each leaflet, as well as from the great central ganglion),
and if the nerve endings are neuro-epithelial cells we
should expect the latter to be numerous and fairly
obvious. The contrary is, however, the case, and in
ordinary sections it is hardly possible to make out many
cells of a different type from the pigment cells already
described.

Are the nerve endings different, then, from those
described by Bernard? The osphradium of the gastro-

poda appears to be homologous with that of the lamelli-

branchiata, and so far as general structure is concerned
there is perfect agreement. List has shown that in
Mytilus there are free nerve endings in this organ, and
the author of this memoir has seen and described the
same structures in Pholas and Pecten. Furthermore,
Gilchrist has described free nerve endings in Aplysia.
The most striking similarity in the microscopic
structure of the organ in lamellibranchs and the whelk is
evident, and the description of the lamellibranch
osphradium might be used for that of the whelk. For

BUCCINUM. ; aon

example, List states that ‘‘ the neurofibrillae split in the
epithelium into several branches, form richly branched
networks, and send some fibrillae through the cuticle to
the outside. Since, however, every fibril does not pass
beyond the cuticle, but often only goes as far as its outer
edge, one can easily get the impression that the
osphradial epithelium was ciliated with very delicate
cilia which have partly fallen away in fixation.’’ This
pseudo-ciliated margin is, then, the cuticle with its
numerous neurofibrillae.

Bernard merely states that there is an inter-epli-
thelial network of nerve fibrils, and leaves one to assume
that these are all connected with neuro-epithelial cells.
The real case is as follows:—Below the epithelium runs
the nerve, and with the nerve fibres are multipolar
ganglion cells. From these multipolar cells delicate
neurofibrillae pass out and enter the epithelial region.
Here they branch in all directions and finally reach the
surface. They run through the cuticular seam in a
parallel manner perpendicular to the plane of the epi-
thelium and appear, therefore, as striae in sections.

So far, Ranvier’s gold methods have not been used,
and Bernard’s neuro-epithelial cells have not been
actually re-investigated by the methods he used. The
nerve network with the free nerve endings is, however,
present and is without doubt the important sensory
structure in the organ. By ordinary methods and
macerations no obvious or numerous sense cells were
found. The structure is, in my opinion, identical with
that of the lamellibranch osphradium.

Little need be said about the non-sensory epithelium
of the leaflets. The ciliated band (fig. 45, Osp. cil.) is
about ten cells broad, the cells differ from those of the
sensory area in being compact, more cylindrical, and with

334 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

deeper-staining contents. Bernard is in error in saying
they possess no pigment.

Underlying the epithelium of the osphradial leaflets
is a delicate basement membrane and a connective tissue
layer with prominent muscle fibres. The centre of the
leaflet is occupied by irregular blood spaces, which com-
municate with a very definite sinus running down the -
external margin.

The nerves of the osphradial leaflets are derived from
a central ganglion, situated along the axis of the organ.
This ganglion consists of a central mass of fibrillae, the
neuropil, with the usual cortex of ganglion cells. The
ganglion cells occur, therefore, on all sides, but are,
nevertheless, more concentrated laterally, except where
nerves are given off to the leaflets.

To each leaflet a nerve is given off, and this leaves
the side of the ganglion exactly halfway down. This
principal leaflet nerve runs outwards across the leaflet,
nearer the inferior border than the opposite and attached
edge (fig. 44, Os. n.). It gives off numerous branches on
both sides, but particularly on that towards the mantle.
These latter branches are very regularly arranged, and
from them arise the neuro-fibrillae which enter the
epithelium to form the nerve-net already described.

Tur OTOCYSTS.

The otocysts are spherical sacs formed of a delicate
epithelium. The cells are low and irregular in shape
with but little contents. Two types are present, sense
cells and non-sensory cells.

The depth of cells is only », of the diameter of the
otocyst. The large cavity is filled with calcareous matter
forming an otolith.

—

BUCCINUM. 335
THE RENAL ORGAN.

The renal organ, the so-called ‘‘ nephridium,’’ is
visible at a glance when the shell is removed. It
occupies a position immediately behind the pallial cavity,
on the right side of the pericardium above the digestive
gland and rectum. The renal organ of Buccinwm is not
only one of the most highly developed in the Proso-
branchiata, but one of the most complicated in the
Mollusca. Fundamentally, it is a sac communicating
with the pallial cavity and the pericardium. The wall
of this sac performs a special function—that of renal
excretion, but under no circumstances does the blood
system open to the external world in this organ.
Externally the sac is covered by the general integument
of the body. The outer and inner walls of the renal
organ, that is to say, the former underlying the integu-
ment and the latter resting upon the digestive gland, are
entirely different in appearance. The epithelium of the |
outer wall is prolonged into filaments or processes of
various shapes, which are held together in such a way as
to form ridges projecting into and occluding the lumen
of the organ (fig. 49). In addition to this epithelium,
which is the true glandular layer, there is a supporting
connective tissue layer. The other walls are merely a
delicate transparent epithelium. The renal organ should
be opened by making an incision along the right side
close to the digestive gland. The internal surfaces can
then be examined.

It will be noticed, however, without any dissection
that another structure is present, lying close to the
glandular renal organ proper and forming part of the
walls of the sac. This forms a band about } inch wide
between the pericardium and the renal tissue. It differs

336 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

from the latter in texture and also somewhat in colour.
It is to a large extent quite independent of the renal
organ and has been termed the Nephridial Gland (fig. 49,
neph. gl., fig. 50). Seen from the cavity of the
renal-organ, the nephridial gland differs altogether from
the glandular wall of the former. Instead of ridges
formed of numerous conical and other filaments, the
surface appears as if it were made up of branched and
anastomosing fibres, which generally run transversely to
the long axis of the gland and renal organ.

If a solution of methylene green or séuresfuchsin is
injected into the foot, one finds after a few hours that
the renal organ is taking up the substance from the blood
and is deeply coloured by the dye. The nephridial
eland, however, is quite free from stain. A second
feature of importance is that the nephridial gland is
‘nterpolated into the circulation between the renal organ
and the auricle, and any injection mass forced into the
efferent branchial vessel passes very easily into this gland
after filling the auricle. In fact, the nephridial anne
is a large spongy blood lacuna.

The Renal Aperture.
renal organ is situated on the anterior wall separating the

The external opening of the

gland from the pallial cavity. It hes slightly above and

to the left of the rectum. It is a fairly conspicuous
opening, slit-like, and of about 3 mm. in length. The
long axis of the opening runs dorso-ventrally. The lips
of the aperture are thickened owing to the development
of muscle fibres, which form a sphincter muscle. The

_ opening leads directly into the lumen of the renal organ.,.

which is only separated from the pallial cavity ey the
membrane above mentioned.

The Reno-pericardial Aperture.—This opening is not
situated at the most posterior part of the gland but

BUCCINUM. Say

laterally, on the inner wall of the pericardium, and some
distance away from the main cavity of the renal organ.
The opening is much smaller than the external renal
aperture and often difficult to find, though it is rendered
more conspicuous by the somewhat white lips standing
out on the darker background of membrane. It is not
easy to see what may be the use of the reno-pericardial
canal and opening. In those molluscs in which a
pericardial gland is present as an accessory excretory
organ, the products, of course, would pass to the exterior
by this canal. Pericardial glands are, however, not
widely distributed in the gastropoda, and appear entirely
absent in the whelk.

mammeture of the Renal Organ.

In considering the vascular system of the whelk,
reference has already been made to the large sinus which
les under the floor of the renal organ. Irom this sinus
a number of vessels arise, which, after crossing the lumen
of the nephridium, give rise to branches which pass to
the filaments of the ridges (fig. 49, ren. r.). Hach fila-
ment is an evagination of renal epithelium, and each
contains a blood cavity. The cells of this epithelium are
of two kinds, glandular and ciliated, but the former are
by far the most numerous. The ciliated cells occur on the
summits of the filaments (fig. 52, ren. cil.) and pass
gradually into the gland cells. The gland cells are
regular columnar cells, the height not greatly exceeding
the width. The cytoplasm is very regularly vacuolated,
and remains as a kind of fine network in the cells, staining
reddish violet with methyl-blue-eosin. Very large
vacuoles are not usually present. The nucleus is situated
at the base of the cell. Underlying both kinds of cells is
the continuous supporting membrane of connective tissue

338 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

in which occur scattered muscle fibres (fig. 52). The
blood spaces are extremely narrow but can be traced
very easily after staining with methyl-blue-eosin (Mann).
Every now and then one meets with sections of the vessels
which occur on the ridges (see vascular system for further |
details of circulation in renal organ.) The ridges con-
sidered above belong to what has been termed by Perrier
the ‘* principal system.”’

Running round the left side of the renal organ
proper, between it and the nephridial gland, is a mem-
branous flap ending in a smooth transparent edge (figs.
49 and 51, ren. lam.). It is continued, though very
much smaller in size, along the posterior margin. From
this lamella a number of secondary lamellae arise and run
at right angles attached to the wall of the renal organ
between the ridges already discussed. These little
partitions (fig. 51, lam. pr.) are hidden by the ridges,
but can be seen quite distinctly if the latter are just
pulled aside. In order to examine the structures further
it will be necessary to macerate the gland—and to remove
the glandular filaments entirely.

It will then be seen that the secondary lamellae
give off in their turn tertiary lateral lamellae,
which finally divide up into very delicate branches
(fig. 51, lam. S.). This secondary system does not
appear to play any great part in the réle of
excretion. It seems to be confined to the highest
prosobranchs, and Perrier regards the two systems as
equivalent to the two lobes of the renal organ of the
Volutidae. The histological structure of these lamellae
is very different from that of the filaments. In section
they are of considerable thickness. There is the same
bounding layer of glandular cells, but they are more
cubical than columnar. The greater part of the structure

BUCCINUM. 339

is filled by a mass of cells whose boundaries and nuclei
are often quite difficult to make out (fig. 53). The reason
is that the cells are filled with refringent granules (fig. 53,
C. gr.), possessing great affinity for eosin stains. These
granules when stained mask the nuclei. Under low
powers of the microscope the substance of the lamellae
looks therefore like a granular or even fibrous mass of
connective tissue. In places, cells occur possessing none
of these peculiar contents, and as a result cell boundaries
and nuclei are easily determined, and the whole looks like
an island of cells (fig. 53, C. 7.) in the midst of the dense
staining mass. In many cases it actually appears as if
the granules were first developed in the nuclei.

Finally, one finds blood lacunae occurring amidst
the mass of cells as in the filaments.

NEPHRIDIAL GLAND.

The structure of the nephridial gland is exceedingly
interesting. It is composed of a somewhat compact
lymphoid tissue, a fibrous groundwork with numerous
lymph cells. Everywhere blood lacunae are to be seen.
The most striking feature, however, is the presence of
long canals, evaginations of the wall of the renal organ,
which extend into the nephridial gland and end blindly.
Sometimes the canals branch slightly before terminating.
The canals are about ;4, mm. in diameter and are lined by
ciliated cells. There is no trace of any opening between
the nephridial gland and the lumen of the renal organ.
The function of the nephridial gland does not appear to
have been satisfactorily ascertained. This kind of state-
ment recurs too frequently in descriptions of invertebrate
structures. The injections made in the course of this
work give only the negative result that the gland is not
excretory like the renal organ. The experiments were,

340 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCTETY.

however, neither numerous nor detailed enough to allow
of a really definite statement being made. On the other
hand, the intimate relation between the nephridial gland.
and the vascular system implies some function connected
with the composition of the blood. Perrier discusses two.
theories, the first that it may be an organ for reserve
matter, the second that it is concerned in the formation of
blood. The author of this memoir is inclined to believe
that the gland is a lymph organ with some additional
function: further work is being carried out on the
subject.

The morphological value of this gland is another
point of considerable interest, particularly in connection
with the attempts made to discover a homologue in the
Monotocardia of the second renal organ of the Dioto-
cardia, or to determine which renal organ, if any, has gone
from the Monotocardia. Originally it was supposed that
the renal organ present in the whelk represented the right
nephridium of the Diotocardia. Perrier took this view,
and starting from the fact that in Patella both organs are
present and both are situated to the right of the peri-
cardium, he pointed out the resemblance in position of
the nephridial gland and renal organ in such forms as
Buccinum. His theory, in short, was that the two renal
cavities of the lower gastropods had fused, and the left
renal organ had passed to the right of the pericardium.
Then the right organ had kept its true primary function
of excretion, whilst the left became the nephridial gland
by development in its walls of tissue with a new function.
As further proof of this view, the intermediate forms—
Haliotis, Turbo, and Trochus—are brought forward, in
which, whilst both renal organs are still present, the left
has almost lost its function of excretion.

Against this we have the fact that the renal aperture

> BUCCINUM. 841

of the Monotocardia lies to the left side of the rectum, and
embryological evidence supports strongly the view that
the *‘kidney’’ of the whelk represents the left renal
organ of the Diotocardia. This conclusion seems at
present to be the most probable, and though in Patella it —
is the left kidney that is reduced, the Docoglossa can
hardly be considered as on or near the line of evolution
of the Monotocardia.

REPRODUCTIVE ORGANS.

Under this heading will be considered:—(1) the
Gonads, (2) the Gonoducts, and (3) the organs of
copulation. The sexes are separate in Buccinum, as in
most prosobranchiate molluscs, and the difference is
well marked externally owing to the male possessing a
large penis which lies folded back in the mantle cavity.
The gonoducts are well differentiated and quite separate
from the renal organ, having their own openings to the
exterior.

The Male Gonad (fig. 54, Go.), which resembles
the female in shape, size and appearance generally, is
situated upon the digestive gland and in close contact
with it. Both are in fact covered by the same membrane
forming the integument. ‘The testis extends from the
cleft marking the point of contact of the two lobes of the
digestive gland, along the right side up to the very tip
of the spire (fig. 54). By reason of its more or less bright
yellow colour it stands out markedly against the dark
coloured tissue of the digestive gland. Slightly ventral
to the gonad on the right side, and on the surface of the
so-called “‘ liver,’’ so that it is visible to the eye without
any dissection whatever, runs the male duct, the vas
deferens (fig. 54, V. def.). It is at first very narrow, but
gradually increases in thickness as it approaches the

;

849 TRANSACHIONS LIVERPOOL BIOLOGICAL SOCIETY.

region of the stomach. During all this length it receives
branches from the gonad and is coiled and folded in a very
characteristic manner. These convolutions become
more complex as the vas deferens passes forwards, and
finally it forms a tubular storing chamber in the region
between gonad and renal organ (fig. 54, V. def.’). The
actual length of the vas deferens is several times the
distance from the tip of the spire to the end of that duct
in a straight line. After the convolutions the vas
deferens becomes again somewhat reduced in diameter
and, with fewer folds, passes forwards towards the
thickened body wall of the “‘ neck’ region. Just upon
reaching the pallial cavity, it turns in a spiral manner—
this part being again of somewhat greater diameter—and
then runs directly to the base of the penis under the body
wall, which forms the floor of the body cavity.

The Penis is a permanent organ of very considerable —
size, which arises somewhat on the right side of the neck
(fig. 54, Pen.). It is spatula-shaped, flattened, and
broadest at the distal extremity. Though capable of some
contraction, this tough muscular extension of the body is —
so long that it would protrude from the shell were it not
for the fact that normally it is bent back and lies hidden

in the mantle cavity. A little distance from the end

there is a small tentacle at the apex of which the vas
deferens opens. The vas deferens, however, does not run
directly through the penis. It winds on itself in a spiral
fashion, running through a tunnel from the base of the
organ to its aperture on the tentacle. The male organs,
it will be seen, are comparatively simple, and no
accessory structures are connected with the vas deferens.
In some gastropods the penis has been found to undergo
seasonal variation in size—no trace of this has been
noticed in Buccinum.

i ee a a ee

BUCCINUM. 343

Sections through the testis show a_ tubular
structure, the tubes opening into one another until
finally a small duct is formed which joins others and
makes up the canal opening into the vas deferens.
In the ripe testis the germinal epithelium, which forms
the wall of these tubes, gives rise to spermatocytes, which
break away and fill up the marginal area of the cavity
(fig. 56). Nearer the centre of the cavity are numerous
smaller cells, the products of division of the spermato-
cytes, and themselves the spermatids (fig. 56, Sptd.).
Both spermatocytes and spermatids are characterised by
the small quantity of cytoplasm and the densely-stained
nuclei. Complete series of stages in spermatogenesis
may thus be seen, but the bodies formed are not of such
a size as to make observation easy. The nucleus of
the spermatid gradually elongates and becomes more
attenuated to form the head of the spermatozoon.

Retzius was the first to discover two kinds of
spermatozoa in the whelk. The presence of two kinds of
spermatozoa in the gastropoda was noticed so far back as
1875 by Schenk in Jfurex. Since then the double
character has been recognised in several species of
Prosobranchs. The two different forms have been

’]

designated by the names “‘ hair-like’’ and ‘‘ worm-like.”’
The former are long fibres with an extremely long head,
almost half the length of the sperm (fig. 57, b.). In
ordinary stained preparations these will be probably the
only sperms recognised, and the only differentiation will
be into the long narrow head and tail. At the anterior
end of the head is a small, clear, vesicular cap. From
this point a central fibre is visible running down to the
base of the head. Very often in fixation this contracts
so as to throw the head into a spiral. A middle piece
follows the head, but is of the same diameter. The tail

344 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

is also of the same diameter and retains this until the

- pointed extremity is reached.

The ‘‘ worm-like’’ spermatozoa (fig. 57, c.) are
rather wider and shorter than the “‘ hair-like,’’? and the
region of greatest diameter is near the anterior end. No
differentiation can be made out except that the cytoplasm
possesses a number of granules. Retzius states, however,
that in macerated preparations a long fibre can be seen
running down one side from end to end.

The vas deferens is lined by ciliated cells, and in the
reproducing season is filled with spermatozoa, any cut
causing a white milk-like fluid to ooze out.

The Female Gonad occupies the same position as that
of the male, and running along its right and ventral side
is the oviduct, which lies in very much the same position |
as the vas deferens in the male, and begins by being an
extremely small canal. It is very different, however,
from the male duct in being perfectly straight without
the peculiar windings (fig. 55, Ovd.'), which seem to be
characteristic of the male gonoducts. One meets with the
same convoluted vas deferens in other groups of animals.
It is probably due to the fact that development of the
spermatozoa takes place after the spermatocytes have
been cut off from the epithelium, and also to the absence

of any large storing organ for the spermatozoa. The

oviduct is a narrow tube with delicate walls. In fact,. it
resembles a large blood vessel, running along the right
side of the digestive gland close to the surface (fig. 55,
Ovd.'). Ata point below the renal organ,.it turns abruptly
at right angles, and immediately takes on a very different
structure (fig. 55, Ovd."). The walls at once increase in
thickness until the tube forms a cylinder about 4 an inch
in diameter. The walls of this are, right and left, about
+ inch thick, and the lumen in section is a long crescentic

= ->-

BUCCINUM. 345

sht running dorso-ventrally. These walls are perfectly
white and of a very peculiar cheese-like consistency.
This posterior glandular portion of the oviduct, after its
abrupt beginning, takes again the anterior direction, and
runs along to the right of the rectum, the rectum, in fact,
lying upon it (fig. 59). It eventually opens by a small
orifice into the pallial cavity.

The ovary is also tubular in section, the wide tubules
of which it is composed being arranged at right angles
to the surface of the gonad. In a hand section the
parallel arrangement of ovarian tubules may be
distinguished quite easily with the naked eye. In
transverse section they are five or six-sided (fig. 60),
and are composed of a thin wall of germinal epithelium
(fig. 60, Ge. ep.)—flattened cells—which here and there
gives rise to large egg cells. These contain a conspicuous
nucleus with nucleolus. One interesting fact is that the
germinal epithelium gives rise to delicate follicles of
flattened cells, which surround the developing eggs
(fig. 60, Foll. ep.). This is seen again in the
Cephalopoda, but does not seem to be the rule in the
Lamellibranchiata. It is certainly not so in Pecten or
Cyclas, where the stalked egg projects freely into the
cavity, surrounded only by the egg membrane.

EMBRYOLOGY.

Fertilisation takes place internally, but details of
the exact procedure have so far not been observed. The
eggs, surrounded by a transparent viscous mass of
albumen, are laid in capsules which are deposited in
considerable numbers attached to each other, and often
fixed to floating objects or to molluse shells, rocks,
crustacea, etc., on the sea bottom. Whelks’ egg-capsules

346 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

form one of the most familiar objects of the flotsam and
jetsam thrown up on our shores.

The individual capsules are flattened on the under
side, and when seen from above they are oval in shape.

A clearer idea of their appearance can be best obtained

by reference to the figures. The wall is made of two
layers of membrane, both of which are perfectly smooth
on the ventral flattened side (fig. 61). The outer wall
appears crinkled on the convex surface (fig. 65). This is
due to the presence of delicate fibres, which, however,
are of short length and often branch.

The outer walls of adjacent capsules are continuous,
so as to join the yellow capsules together in masses.

The eggs of the gastropoda are not always laid in
chitinous capsules of this kind, there being other modes,
1.e. (@) free without any protecting capsule, (6) with
calcified membrane, (c) deposited in ribbons (Opistho-
branchs), and (d) in gelatinous masses. The capsules
of the whelk have been supposed by most zoologists
to be formed by the oviduct, and the most distal or
uterine portion of that tube is extremely glandular.
Cunningham, however, in 1899, in a letter to “* Nature,’”’
announced that the egg capsules of Buccinwm and Murex
were formed by a secretion from the anterior groove of
the foot. The eggs must therefore pass round from the
pallial cavity, imbedded in a quantity of gelatinous
material, to the under surface of the foot. Then in the
groove they become surrounded by the chitinous capsule.
Cunningham’s discovery has apparently never been
noticed, and no information on the subject appears in
the textbooks. Not only does Buccinum torm its egg

capsules in this way, but the same thing apphes to

Purpura, and Pelseneer has actually found the capsules
in the ventral pedal gland.

BUCCINUM. 347

The exact number of eggs placed in the capsules also
seems to have been overlooked by most workers. Koren
and Danielssen, who give the largest numbers, say that
variations from 6 to 800 occur in Buccinum undatum.
Our counts have given from 49 up to 2,419! Moreover,
the high numbers were by far the most common. Few
capsules contained less than several hundred eggs. In
what must be regarded as a very small clump of egg
capsules (about 2 cub. inches) we calculated that there
must be about 200,000 eggs.

It is a well-known fact that the number of young
whelks leaving the capsules is very much smaller than
the number of eggs placed therein. Here again, the
figures vary, but the average 1s probably somewhere
around the figure 10. Each capsule is penetrated by a
small oval hole, which appears always in the same place,
near one margin of the flat side (fig. 61).

The eggs are 0°25 mm. in diameter, the young shells
at the time of leaving the capsules are 3 mm. from the
tip of the spire to the end of the siphon, and three
whorls are present of rapidly increasing size.

The embryology of the whelk must be left for a
separate work, any consideration of it in a detailed
manner would be beyond the usual limits of a memoir of
this kind. Koren and Danielssen, in the ‘* Fauna
littoralis Norvegiae,’’ were probably the first to describe
the sequence of events taking place in the egg capsules.
Their account is extremely interesting, though the
treatment was naturally limited by lack of the methods
now available for embryological research. The eggs
when laid are imbedded in a perfectly transparent
viscous mass. As development takes place, this albumen
becomes less and less viscous, and the eggs become
crowded together. Now, according to Koren and

348 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Danielssen, little groups of eggs, containing from 30-
60, sometimes even 180 individuals, separate out, and
around them a delicate granular mass appears—a
supposed exudation from the eggs. Each little group of
eggs becomes an embryo, and the granular mass takes on
the form of a definite limiting membrane, which gives
later the shape of the larva. :

What really happens, however, is the much more
probable sequence observed first by Carpenter in
Purpura lapillus. According to this investigator, only
a few of the eggs in a capsule are to be looked upon as
true ova. The remainder are termed ‘‘ yolk spherules.”’
The distinction between the two is manifested at the time
of the first segmentation, the yolk spherules dividing
into two equal hemispheres, the real ova into a larger
and smaller segment. Segmentation of both takes place,
however, and it should be interesting to determine
whether this is due to an actual difference in the eggs laid
or to fertilisation by the different kinds of spermatozoa
now known to exist. As the embryos develop they
commence actually to swallow the segmented yolk
spherules, and it is these carnivorous embryos which
Koren and Danielssen took to be clusters of eggs with an
exuded membrane.

Bobretzky describes in some greater detail the
embryology of a gastropod supposed to be Fusus (sp. ?)
Whilst, however, he gives in his account a description
of the formation of blastula, gastrula, and early
embryonic stages, he does not make any mention of the
carnivorous act, and his egg capsules only contained
7-20 eggs! The egg divides into two similar halves,
and then into four large cells by a second division.
These four large cells cut off four small ones which lie
close together. Four other small cells follow these, being

BUCCINUM. 349

cut off in a similar manner, and then by division of the
first four small cells a total number of twelve is reached.
By the segmentation of the small cells themselves, and
by the addition of others a layer of cells (ectoderm) is
formed which gradually grows round the egg until a
space is left at the opposite pole. Here the four large
cells remain visible and form part of the wall of what
may be termed a blastula. The gastrula is formed by the
depression or invagination (very slight) of the large cells,
and the growth of the ectodern at the blastopore. The
further details of this development do not actually apply
to Buccinum. This, however, gives one a hint of the
processes by which the early carnivorous stage is formed.
As soon as stomach and oesophagus are developed, the
embryo starts devouring the eggs in the capsule.

The ectodern is ciliated, and outlines of head and
foot are soon to be observed. The otocysts are visible
very early, and are closely followed by the eyes. At this
stage the embryo is quite symmetrical. The stage shown
in fig. 64 on Pl. VIII, represents a fully developed
embryo some time before the shell and body attains the
size which may be termed the “‘ young whelk’’ stage, at.
which it leaves the capsule. At this period the head is
well developed, and eyes and tentacles are obvious
structures. The velum (fig. 64), in the form of a bilobed
structure fringed with particularly large cilia, has
reached its maximum development, and by that means
the larva moves actively round in the capsule. The foot
(fig. 64) is well developed—the pedal groove (Ped. gl.)
being relatively large, and the otocysts can still be seen,
though they have sunk away from the epithelium and lie
quite deep in the body. The larval shell is also
conspicuous. It is a very delicate chitin-like structure,
not yet coiled in a spiral, and marked by delicate

i
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|
4
t

350 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

longitudinal ridges. One very curious organ must also be
mentioned. Protruding slightly from the pallial cavity
is a vesicle (fig. 64, Pul.), which is apparently an
evagination from the floor of the cavity. This vesicle
contracts rhythmically. It has been noticed in other
larval gastropods and termed the “ larval heart.’’ It is
not, however, to be regarded as an early stage in the
development of the true heart, which is situated much.
further back. .

Further development results in the absorption of the
velum, increase in size, and spiral coiling of the now
asymmetrical shell. Of the thousand eggs that might
have been in the capsule, perhaps ten may eventually
reach the stage, of which the shell is figured on Pl. VIII.
This reduction is due in the first place to the small
number of eggs that can develop into young embryos, and
secondly to the cannibalism exhibited by those same
embryos practically as soon as the first rudiments of an

alimentary canal are developed.

. \\ \\ \ \\
NS NS . aN Ww \ \ :
I YOY MD Ny. ef
So : : ‘ ” \\ A

\) \\ \\\ AN \ \\ \ I\y :

q oo Ky

—_- ~~
2

BUCCINUM. ool

APPENDIX

DISTRIBUTION AND ECONOMICS.

The genus Buccinum is widely distributed in Arctic,
Antarctic and Temperate zones. The species Buccinum
undatum, Linn., occurs all around our coasts, from low-
water mark to 100 fathoms, and over a considerable area
extending from the Atlantic coast of North America to
the Siberian seas. The genus appeared first in the
Jurassic rocks, the species B. undatum occurring in the
Coralline Crag, since when it seems to have become
increasingly abundant in our seas. The Common
Whelk, B. undatum, is the most abundant species and
the most convenient to examine as a type of the genus.
It inhabits different kinds of ground, and several marked
varieties are to be found from the littoral zone to
considerable depths. There seems, however, to be little
agreement in regard to, or scientific classification of,
the varieties.

Whelks are used at many of the fishing ports along
the coast as bait for the long lines. They are caught by
the Manx fishermen from banks of a shelly nature, about
17 to 20 fathoms deep, often near beds of the scallop
Pecten opercularis.. The method employed is to sink
wicker baskets (crab or whelk pots) of the kind used for
capturing the edible crab, baiting them usually with the
latter animal. The crabs are used fresh, and are strung

352 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

on a piece of cane which is inserted in the basket. An
average catch at Port Erin is about 20 to 25 whelks per
two days.

The whelk is also used as food, and large quantities
are usually exposed for sale in London. It does not
appear to be eaten much in the North of England, but a
few specimens are occasionally seen for sale in Liverpool.
In the Isle of Man, and Scotland, the whelks are
commonly termed ‘* buckies’’; in Heligoland ““coxen,”
and in the days of British rule, the English fishermen
who called there were known as ‘‘ coxen clappers,’’ from
their habit of breaking up the shells to obtain the whelk
for bait.

Joubin states* that Buccinwm undatum is very
common on the West Coast of France, and that in the
Syndicat of Portbail a fisherman can collect as many as
200 at one low tide. These he can sell at 1d. to 3d. per
dozen. They are consumed in the country, and not
exported.

A considerable demand for whelks must, however,
still exist in this country, and one finds quotations
regularly in the Mish Trades Gazette, though, unfor-
tunately, the quantities sold or brought in are not
indicated. The prices at Billingsgate appear to vary
from about 4s. to 10s. a bushel. Government reports
give little information, whelks being tabulated with other
edible animals as ‘‘ unclassified shell-fish,’? in the
statistics published by the Board of Agriculture and
Fisheries; but Mr. C. E. Fryer, I.8.0., of the Fisheries
Department of the Board, has very kindly suppled Prof.
Herdman with the following particulars collected for
last year :—

* Bull. de L’Institut Oceanograph. Monaco. No. 218, July, 1911.

BUCCINUM. 315133

QUANTITY AND VALUE OF WHELKS RETURNED AS LANDED
AT STATIONS IN ENGLAND AND WALES IN 1911.

Station. Quantity Value.

East Coast— cwts. Bg
: SL ere 1,292 300
: UL 222 5203. Ce eee eas
: Se ne 12-758 ss oS
| Sheringham .........ccc. S074. TOR ital
eo 4,482 2E 1868
Ipmiebtlinesea .............4.- Qe ee ol
| Dac OS ea acre Side 20 144
0 TESS) 0 TORO 22: | eee
q Ramsgate ........ SN eee BAG o>: sects eeeeleges
| SGT s 2 let Lee eee eee 43,142 8,238

South Coast—

“Toc cet \. an AO 54 149

“C1 ee 686.5. -.23: 171
| TMM EGHEOUIGI 003.6. csv ccenss es T74 115
SOMPMATUPTON nec cssesseses- 337 165

| NOSE Lane ae pid +: Sie ne 600

MEW Sta vLOUS -,. 5.46. ..0- 1 Oe Saeeey 212

MGIRAT, P28 Sera this’: « AGB,450) . ... ) WOghkO

The returns and remarks in trade journals with
reference to Billingsgate are reflected in the statements
made in the Annual Reports of the Inspectors of
Fisheries for England and Wales. Thus one finds that
practically the only ports where whelk-fishing is carried
on are situated between Grimsby and Southampton.
This probably means that the fishery is governed by the
presence of a large market for the mollusc in the south-
eastern area of England.

354 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The following official statement shows the small
extent of the fishery in the Lancashire and Western Sea
Fisheries District :—

Landed at Liverpool—

Wt. in ewts. Value in £’s.

OOO. eerste HG, Beene orc eae nil.
2) epee a ee D CWS. «deco se ee nil.
U9O8: Gee oo: A CWHSE wo c0 iste B

L909." ees de ee: WE 4 gy ee nese ake nil
HS es eich Oe Lewte ..Ac0peuee nil
LD Bee 8 Pa Pe el peices REE nil.

As a matter of fact, however, the real value of the
fishery must be higher than is represented by these
figures. It is known that four to five hundred whelks
are landed weekly by small sailing boats, and these sell
at ls. to ls. dd. per hundred. In addition to these,
others are apparently landed by longshore fishermen and
find their way with illegal-sized flat fish into some of the
smaller fish-shops. :

In addition to whelk-catching by means of wicker
pots, dredges are occasionally used; ‘“‘trotting’’ is
another method adopted in the south-eastern districts of
England. A number of shore crabs are strung together
with a needle and string, so as to make a bunch. These
are sunk to the bottom and left for a time; they are
afterwards drawn up and the whelks removed from them.

In England, some years back, the trade in whelks
must have been of considerable importance, for one finds
that the Lynn fishery alone supplied about 1,250 tons a
year, for which about £10,000 was paid, and Grimsby
exceeded this with a value of £22,000. These figures
probably include Fusus antiquus (the ‘‘ hard whelk’’),

which seems to have been more prized in some markets.

BUCCINUM. 355

The name Buccinum comes from “ buccina,’’ a
trumpet, but it is difficult to get evidence that Buccinum
undatum was ever used as a musical instrument. Species
of Triton seem to be most used for this purpose, and
Turbinella (the Chank Shell) is used in Ceylon by the
Buddhists. The Chinese use a species of Fusus for the
same purpose. The shell of Pusus antiquus is also used
occasionally in the Scottish Isles as a rude lamp, in which
to burn the oil of sea-birds.

The two closely related families, the Muricidae and
Buccinidae, contain various genera and species, which
have figured, perhaps, more than most marine animals,
in the histories and traditions of ancient peoples. They
have played an important part in religion, mythology
and war, in the production of ornaments, and in the
preparation of the famous ‘‘ Tyrian’’ purple dyes.

The whelk crawls about the sea-bottom by means of
its muscular foot, and when kept in large aquarium
tanks occasionally creeps above the level of the water.
It seems to remain, however, about a foot or so above the
surface, and never crawls further up. It was observed
at Port Erin, on every occasion on which whelks were
turned into a large tank into which a stream of water
was running, that they found their way towards the
entrance pipe. ‘They then remained either just below or
just above the level of the water, in the latter case bathed
by the spray, although the water was well aerated in
every part of the tank. |

The whelk appears to be omnivorous so far as its
diet is concerned, and dead (but fresh) or living
organisms seem equally acceptable. The food, which is
scraped away by its jaw apparatus, is taken into the
stomach in a finely divided state, and little information
as to its character can be gathered from an examination
of the stomach contents. —

356 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The fishermen at the Isle of Man have found that
the best bait for their whelk-pots is Cancer pagurus, the
edible crab, used fresh. The whelk inserts the proboscis
through the holes broken into the carapace of the crab.
Whelks have been observed to attack living lamelli--
branchs, and on one occasion at Port Erin a large
specimen of Buccinum was observed to prevent a Pecten
maximus from closing its valves by inserting the anterior
end of the shell between them. It then attacked the
adductor muscle with its long proboscis, and so at the
same time obtained food and disabled the closing
mechanism of the scallop. The same process of disabling
the prey has been observed by Colton in the American
genera Pulgur and Sycotypus. Colton experimented with
these animals, and found that when an oyster was given
to a hungry Sycotypus, the latter crawled on the top of
the oyster and waited until the valves were opened. It
then rotated on the columella and inserted the end of
its own shell between the valves. Forty minutes later it
left an empty shell. Fulgur actually hammers the
margin of lamellibranch shells by grasping with its foot
and contracting the columellar muscle sharply. The
proboscis is then inserted into the orifice so made. The
impression one usually derives from the literature of the
subject is that the whelk actively bores through
lamellibranch shells by means of its odontophore.
Colton states, however, that the radular teeth of Pulgur,
Nassa, Lunatia and Purpura do not appear as if worn
down against a hard substance, but broken off irregularly.
To this lst the whelk can be added, for the old teeth
present a most ragged appearance (fig. 15).

On one occasion a whelk was observed attacking a
dead Nephrops (the Norway lobster). It held the
crustacean by means of the anterior part of the foot,

BUCCINUM. ob

using the latter to envelop the posterior part of the
abdomen of its prey. The radula was then brought into
action, and a hole was bored through the chitinous
exoskeleton until the proboscis could reach the muscles.
Here again the use of the foot as a grasping organ may
be noticed. In its turn the whelk must fall a prey to
many other inhabitants of the sea, and one often wonders
what has happened to the former owners of the numerous
whelk shells now found occupied by hermit crabs (see
Text-fig. on p. 350).

The cod feeds to a certain extent upon the whelk,
though this is by no means its chief food. Remains
of the whelk have also been found in the stomach
of the dog-fish. Curiously enough, in most cases these
remains include only the fleshy part of the animal
and the operculum. This implies that, contrary to
expectation, the fish either bites off the protruding part
of the whelk or otherwise achieves the apparently
impossible in removing the animal from its shell.

Quite recently, Dr. C.G. Joh. Petersen has published
a very interesting paper on the relations of the whelk to
the fisheries of Denmark. In 1909 a committee was
formed in that country to consider the harmful animals
of the sea-fisheries; and five animals were black-listed—
the sea scorpion, the stickleback, crabs, starfish, and
whelks. Petersen suggested that before money was spent
on efforts to exterminate these creatures, experiments
should be made to determine the possibility of such a
proceeding. The whelk was selected for the experiments,
and the Board of Agriculture allotted funds for the work.
The harm done by the whelks was known and had been
investigated some time earlier (1895). It had been found
that the whelk attacked the plaice entangled in fishing-
nets. Although unable to seize actively-moving fish,

358 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

they very soon found and attached themselves to the fish
that had been caught. The whelk proceeds by boring a
hole through the skin, inserting its snout, and then
devouring all the muscular tissue, leaving nothing but
skin and bones. The molluse will not touch rotten fish,
but experiments have shown that it will devour anything
so long as it is fresh. Often enough, 10 to 20 whelks
would be found attached to a plaice, and the fishermen
estimated that one-third of the year’s catch was lost in
the region investigated (Thisted Bredning, in Denmark).

The experiments carried out by Petersen were of a
two-fold nature—(1) to determine the number of whelks
in a certain area, and (2) to find out whether these could
be economically exterminated by capture. The first
part of the work was carried out by using an instrument

Ue)

called the ‘‘ bottom-sampler,’’ and also by the employ-
ment of a diver. The latter caught from 106 square
metres 100 whelks, 36 square metres 128 whelks, 106
square metres 70 whelks, 106 square metres 97 whelks.
In the last two cases the man reckoned that he had
only taken one-third of the whelks present. From the
results it was calculated that 130 million whelks were
present in the region (an area of 65 million square
metres). |

One motor boat, with 240 traps, was able to catch
3,8454 bushels (45 tons of whelks!) in the same district,
between April 5th and November 8th.

On the whole, it was concluded that the extermina-
tion by catching would not be an economic method,
especially since there did not seem to be any great use

for the whelks caught in the district. Petersen also

states that six or seven whelks’ opercula may be found in
one cod’s stomach, and comments on the rarity of any
remains of the shells.

i i

———————————=———— TS ee
~

BUCCINUM. 359

PARASITES.

Two extremely interesting parasites have been found
in the whelks from Port Erin. One of these is a
Coccidian (Merocystis khathae, Dakin), and the other is an
endoparasitic Turbellarian first found 14 years ago by
Jameson, and re-discovered this year in whelks from the
same district. The Turbellarian (Graffilla buccinicola)
belongs to a genus of parasitic rhabdocoeles which occurs
frequently in mollusca, but at the time of Jameson’s
discovery Graffilla had not been found in British waters.
No further details of its anatomy will be given here,
because it seems advisable that the species should
undergo a thorough re-examination. One point, however,
must be referred to. Jameson states that the parasite
occurs in the kidney and “kidney duct’’ of Buccinum
undatum and Fusus antiquus, and that almost all whelks
examined were infected. My specimens were examined
fresh, and considerable numbers were investigated. In
all cases the parasites were found in the stomach and
rectum, and there were as many as 14 seen in the stomach
of one individual.

It is somewhat difficult to conjecture what the
‘kidney duct ’’ referred to in Jameson’s description can
be, for the renal organ opens directly at one end into the
mantle cavity. The minute reno-pericardial canal can

hardly be called a kidney duct!

360 TRANSACTIONS LIVERPOOL BIOLOGICAL SOGIETY.

INDEX

PAGE

Alimentary canal 2..2...2.,0-.+.-- 280
histology ... 287

ATS Bexek cote ie eee Zt aon
DENG ig fe Me eee ey Ae et alt ee ee 310, 311
Geplaliciee..cs-: forthe 311, 312
WISCeralll ct eaccducche. SLT sls

Pe\v aires aia OTOGEN “rodeo dic cabal ouadace 312
Pala es eecoa cue. 311

pedal. sees emmiasaccss cot 312
Gembtacwlar, sss scence 313

Auricles pitscctte sc aac eee: 310

Auriculo-ventricular passage ... 310

Billingsgate returns ...........:... 357
Body whorlvol shelly...) .-2--.- 258
Branchial vessel afferent 274, 317

efferent 274, 317

Buccal -aRvenya wee rscnce sc ce teasca 312
CANS MAM. onemacieiaes te oe 320
Carnivorous gastropoda ......... 258
Cephalic simus, eerssersee ase eee 314
Cerebral ganglia ............ 319, 320
Classthteationm.sss.ssne- une 253, 254.
Columella muscle .................. 263
COrm@aincanetvie seat ee so eee 326
Corneal scelllstety jase ste sneer 326
Ctenidiuinay S.niisceeese sree oe 269, 273
CEenicial taxis seen eee ee 274.
Dextral ‘shells ss geccseaenanene ae te 257
Digestive elandiy es rere seen e es 286
structure 293
Distributiony a2. ce seers 351
RICOMOMICS 7 Aeaacts cea oos eee eee 351
Beg capsules(-.c-scaece 269, 345, 346
Kimo bryOlogiy (eases eaecemeee se 345
Enemies of whelk ............... 357
Fisheries, relation to ...... Bol, ool
Fishing, methods of _...... 351, 354
Rood oF wihellcims-. tacceea ede 355
HOO? Gree dnc gaccueoaueee ,. 205, 262
glands anterior | ~).:,0..-20- 263

MEG ATI Sct vere 263

i PAGE,
Gastropoda....;.02..-...emeemee 253, 254
Ganglia buccal! ~....-steseeeeeeeee 320

cerebral {).-..tecter 319, 320
subintestinal ............ 323
supraintestinal ......... 323
pleural ..2.0..2. 3.t-eeeeee 321
pedal .....v..ecseceeeeeeee 319
visceral..:.:0....40-eeeneee 324
Geological distribution ......... 351
Gill, connective tissue of... 276, 277
epithelium of) --sctes-e-sees 27
leaflets (of .....ciescnieeteeneee 273
supporting membrane of 276
Gland, digestive i2s22---eare 286, 293
hypobranchial ...... 270, 278
‘of Leiblein =< asem 284, 288
mantle edge ......... 272
MUCOUS) Scenes 270, 278, 279
nephridial~ “a.-20e8 336, 339
pedal -\..2s5.82-eeseee 262, 268
POISON. \ssaaenekecrnisepenee 289
rectal ..cc.-aseaseeeeneeeee 287
salivary peer 283, 284, 290
Gonads ......5.06;5 esse eee eee 341
Head —a..0.56.scegeeeee eee eee 255
abnormal — S2aesee-ae eee 255
Heart......52.00. sees ateeeete 310
Hypobranchial gland ...... 270, 278
Injecting methods--7.... sss: eeeeee 308
Intestine ~ i... ccusens eee 286, 294.
Intestinal ganglia .................. 323
Lens oie scisesinsceentan eee (ae eeeene 327
Littorina — .... 0.0 +. 1c scemeeeeenee 307
Locomotion ....st-..e--heeaeeeeeees 264
Mantle....:.cdecceane nese 255, 269
CAGE ....00.0000.020 see 271
gland ....::sccseeeeaeee 272
Monopectinate ...............ss+00: 274
Monotocardia ~ .....-csssseeeeeeeene 254
Mucous gland: |... aeeeeeeees 270, 278

BUCCINUM. 361
PAGE PAGE
Nerve centres anterior ............ 318 Fadular teeth. ..c.6..2.--c00-crs-0s 301
aula!) - Se 323, 324 variation in teeth ......... 301
Ott. Sees 319 formation and growth 304, 305
TRL2 (eee 319 RECHUIA. cccvcachcpqteacsuseveomesucrse 287
of proboscis and proboscis Plands = Fa snvaxtacsnnes sees 287
Vie) 321 Renal aperture .............+. 271, 287
MRC EMIN cara on a veneevesaee 325 QPCR wane cess petina teehee 335
endings in osphradium 332,333 blood supply ...... 315
Nervous system................0005 317 principal system... 338
Ue 335 secondary lamellae 338
Nepbridial 2) Seen 339 sinus system ......... 314, 315
Nephridio-cardiac vein...... 310, 316 Reno-mucous vessel ... 309, 314, 316
Reno-pericardial aperture ...... 336
Reproductive organs ............ 341
‘Khynehodaeumy ..<.s6s- s4ssoee 282
OMUODIARED ©: ....-6.---2.002000c00e 304 Phynechostomie:, : 5... sass deeeee 282
Mdontophiore .....-.........+ 283, 295
AMES YE access s:a/0s's- 296
Cartilage .........+0. 297 Salivary GUCts, ..<socas-eeneen 284, 290
histology 302 glands s.s'szsenke- a 290
MPMI CIOS ico cececies vise 296 SENSES. ONCANS : oiisc<sedeponts aeenee 325
mode of action ... 299 POMEL iianss Sagi caspian ceipetestan se eRe eee 256
Werephagis” ........0......++. 283, 290 LOTMAbLON -$...<snhansepeames 261
eCnecum ......... 285, 292 JAVORS:- un ro.2 ev nts cceueeceeeen 259
epithelium ......... 291 sinistral... covstraattneceees 257
MSEC © oo acon sees enue. 256, 265 SUDO. co. cis /eo chsh omer ees 258
5 326 SAMUS ooo. tn ple angen: 257, 258
JS ot 270, 329 SETUCHHIGS <2. «3:12. eee 260
Te BHOL Bs 5-s.-.00-00 330 Silica wus teeth. <i... ake aoa sans 307
nerve endings 332, 333 SPELNIALOZO2) o's 2 ssn esnjnber ean she tae 343
PORES OY Sorcies so vcsiesveo.e.. 320, 334 SS HEIGCLIOED: © (6.05: sous ce Sand eee 328
DMM lates en cs ceusnecrncesseee 345 SLOMAN. 5.08. csanet eae 285, 292
Streptoneura def. ......... 253, 254
Streptoneurous twisting ......... 317
Pallial artery .......02...sseeeee 311
MEME Edina» aeina rnin’ ea oes 269
RCT a gv'Sivp isis vacsescsoce 269 MENT AGIOS s saise'ctisn veer ete neeetee 255
circulation ...... 316 supernumerary ...... 255
OE o.1/05/5 9%, 000 323, 324 Tentacular artery — ...........0.. 313
EADS isiG's sia d3-oves'e nee. 270 NOIVG s. snaierdeeneoseare 320
SEI gasps uses. 0ciensccscree see 359 TRSEIS).. saibhins yas ins secen ahem eee 341
Ee 807, 341 “Triangle lateral ** casesaweeeee 339
Pectimibranchia ...............000 254
(05 319
asntcs sviphvis's sos 262, 268 Vaginal oviduct ..0.....2..s20dve- 345
00 le 256, 267 Vascular SyStem .:.c.ésoncvenreape 307
PUNGU TE) desc yccvecscstannsorscens 342 Venous System: ..narvan tons chaheees 313
ee ive se esesdiviisaes 319 Ventriclo ..drvbncsdinetiosahe ee 310, 311
Periostracum ................ 258, 259 Visceral sorte .<isscencessaces ol), als
secretion of ...... 261 commissure 317, 318, 323
MIN Sdn anccvecscsisorasee 282, 287 PANOUE «<. nanatecceameeee 324
MMTAL PANCIIA ........0cccedeeceree 321 MASS 55 50's ha0kd suepaeahes 255
MEBORCKDOLIC .........0s0ccsenceseess 282 NEPves. . <.cchigacntvanenes 325
MRCIIDOLIC =... ss sacoeceeseriees 282
EE p85 cco snarndepencdiinvens 280
BRORODEATICDIA 26.0 cc0ceisvesceerers 254 WINE 3 «scab cersevanancenasneeenaier 257

EXPLANATION

362 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

OF PLATES

ReFerRence Letrers

buc. = Buccal artery.
c.= Anterior aorta.

cut.’ =Cutaneous artery.
col. =Columella artery.
g. = Gastric artery.

go. =Gonad vessels.

. pall. = Pallial artery.

. ped. = Pedal artery.

pen. = Penis artery.

vis. = Visceral artery.
Af. ct.=Afferent branchial vessel.

DRED DDR RRR

Ao. = Aorta.

Br. v. = Branchial vessels.

bucc. = Buccal ganglia,

c. =Cerebral ganglion.

C. c.=Occlusor muscle.

C. cav.= Blood spaces in oeso-
phagus.

C. p.=Cerebro pedal connective.

c. d. m.=Centro dorsal muscle.

c. gr.=Granular cells of renal
lamella.

Caec. = Oesophageal caecum.

cae. w. =Connective tissue of caec.
wall.

cart. = Cartilage.

cer. com. =Cerebral commissure.

Col. mus. =Columella muscle.

Con. t.=Connect. tiss. gland of
Leiblein.

Con. =Connective tissue.

Cor. = Cornea.
= Ctenidium.

ct. ax. =Ctenidial axis.

ct. con. =Connective tissue of gill
leaflet.

ct. e.’ = Epithelium of gill leaflet.

ct. ep. =Ctenidial epithelium.

ct. gl. =Connect. tissue of etenidial
axis.

ct. mus. =Ctenidial muscles.

ct. n. =Ctenidial nerve.

D.m. s.= Dorsal trans. muscle
sheet.

D. pr. m. = Dorsal protractor
muscles.

Dg.’ = Lobe of stomach.

Dg.c.”” Dg.c.’=Cells of digestive
gland.

Dg. con. =Connective tissue of
gland.

Dg.d. = Opening of post. diveneaees
gland duct.

Dg.d.’ = Opening of ant. dig. gland -
duct.

Dg. gl. = Digestive gland. -

Dg. gr. =Granules of gland cells.

Dg. nuc. =Nucleus of gl. cell.

Eff. ct. = Efferent branchial vessel.

Foot = Foot.
Foll. ep. = Follicular epithelium.

Ge. ep. =Germinal epithelium.
Gl. c.=Gland cells.
Go. = Gonad.

l.r.r. = Dorsal retractor muscles.

L.t.b. = Lateral odontophoral
bands.

Lam. pr.=Lamellae of secondary
system.

Lam. v.= Vein of renal lamella.

Lens = Lens.

Ling. con.=Connective tissue of
odontophore.

Im. g.=Gland of Leiblein.

M.=Mouth.

_m.v.r. = Median ventral muscles.

mem. = Membranous wall of larva.

Mu. gl. = Mucous gland.

Mus. circ. =Circular muscles.

Mus. col. ='Transverse sections of
mus. fibres. ;

N. ax.=Nerve of ctenidial axis.

n.ot.=Otocyst nerve. -

neph. gl. =Nephridial gland.

Nuc. = Nucleus.

Nuc. Int. C.=Nuclei of instersti-
tial cells.

O. cav. = Lumen of cost iaeam

oe. con.=Connective tissue “with
longitud. muscles.

oe. con.’ =Connective tissue.

oes. = Oesophagus.

Op.= Operculum.

Op. n.= Optic nerve,

BUCCINUM.

Os. n.=Osphradial nerve.

Osp. cil=Ciliated cell region of
osphrad. leaflets.

Osp. gl.=Gland cell region of
osphrad. leaflets.

Osp. so.=Sensory cell region of
osphrad. leaflets.

Osph. = Osphradium.

Ot. = Otocyst.

ov.=Ovary (oviduct in figs. 38
and 37).

ova. = Eggs.

ovd. = Oviduct.

ovd.’”’ = Vaginal part of oviduct.

P.= Penis nerve. ~

Pall. = Mantle.

Pall. ep.= Mantle epithelium.
Pail. gl.=Mantle conn. tissue.
pall. n. & n.’’=Pallial nerves.
Ped. g.= Pedal ganglia.

Ped. gl.= Pedal gland.

ped. gr. = Pedal groove.

ped. mus. = Pedal muscles.

ped. n. = Pedal nerves.

Pen. = Penis.

Per. p.= Pericardial cavity.
Ph. = Pharynx.

Pig. = Pigment.

pl. 1.= Left pleural ganglion.
pl. p. = Pleuro-pedal connective.
pl. r.= Right pleural ganglion.
pr.= Processes of retinal cells.
prob. = Proboscis.

prob. m.=Proboscis muscles.
Ps. cor.= Pseudo cornea.

Pul. = Larval heart.

r.’ & r.”’=Retractor muscles
(dorsal).

R. Sin. =Reno-mucous vessel.

Rect. = Rectum.

Ren. = Renal organ.

ren. cil.=Ciliated cells of renal
epithelium.

363

ren. eff.= Vessels of renal organ.
ren. ep. = Renal epithelium.

ren. lam. = Renal lamella.

ren. r.= Renal ridges.

Ret. c. = Retinal cells.

rhyn. = Rhynchodoeum.

Rst. = Rhynchostome.

S.=Stomach.

S. gl. c.=Salivary gland cells.

Sal. d.=Salivary duct.

Sal. gl. =Salivary gland.

Siph. = Pallial siphon.

Siph. n.=Siphon nerves.

Sptd. =Spermatid.

Spte. =Spermatocyte.

Spz. =Spermatozoa.

St. =Stomach.

St. s.=Stiftchen.

sub. int. =Subintestinal ganglion.

sup. int. =Supraintestinal ganglion.

sup. m.=Supporting structure of
ctenidial leaflets.

Tent. = Tentacle.
tent.=Tent. nerve (fig. 42).
tri. lat. = Triangle lateral.

V. def. = Vas deferens.

V. def.’ =Coiled part of vas deferens

V. def.” =Vas deferens in penis.

V. def. c.= Vas deferens canal.

V. e. m.=Ventral muscle sheet.

V. pr. m.= Ventral protractor
muscles.

V. r.=Retractor muscles (dorsal).

v. 7. m. = Ventral retractor muscles.

vas. d. = Vas deferens.

vel. = Velum.

vis. com. = Visceral commissure.

X. con. =Connect. tissue of mantle.

Zyg.= Zygoneuric connection.

364 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Fig.
Fig.

Fig.

“I GO OF

ieee

Prater I.

Shell of Buccinum undatum. x 4

Section of Buccinum shell, showing septa at
apex. x @

View of part of columella of 5th whorl,
showing attachment of columellar muscle.
<3

Surface view of periostracum.

Superior surface of operculum. x 1.

Longitudinal section of shell. x 40.

General view of animal after removal of shell.

xe I

Puate II.

Roof of pallial cavity with organs of pallial
complex. x slightly.

Ventral surface of foot (expanded). x 3.

Transverse section of pedal groove and gland.
xT.

Dissection showing cavity into which proboscis
sheath is retracted and also rhynchocoel in
which proboscis hes. x 1.

. Dissection of alimentary canal. Slightly

reduced.

. Stomach as seen from digestive gland surface.

Shghtly reduced.
Radula with teeth. x 48.
Old teeth. x 64.

Puate ITI.

General view of odontophore, radula and
muscles from above. Slightly magnified.

Ventral end of radula with attached ventral
retractors. Slightly magnified.

ig. 27.
ano.

Fig.

ck,

Lg.

ale

. 22.

. 2A.
. 20.

26.

. 29.
. 30.

ae
82.

33.

BUCCINUM. 365

Odontophoral cartilage (odontophoral tongue)
with ventral muscles indicated through it.
Shghtly magnified.

Odontophore from above before muscles are
laid bare by cutting dorsal transverse
sheath. Slightly magnified.

Odontophoral tongue.

Anterior end of odontophoral cartilage with
ventral protractor muscles.

Transverse section of cartilage and radula
muscles. x 480.

Transverse section near tip of odontophoral
tongue.

Transverse section of pharynx wall. x 200.

Transverse section of oesophagus. x 30.

Puate LV.

Section through salivary gland and duct.
x 450.

Epithelium of oesophagus. x 800.

Transverse section of oesophageal caecum.
x 60.

Transverse section of digestive gland.
x 200.

Section through wall of Gland of Leiblein.
x 450.

Transverse section through two gill leaflets.

Transverse section through gill leaflet more

distal to fig. 31.

Transverse section through gill leaflet more
distal to fig. 32.

(The three sections are consecutive and all x 100.)
Fig. 34. Cells from inner surface of wall of gill. x 800.

366.

ig. 39.
. 40.
WAL:
. 42.

ig. 50.

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

oO.
. 36.
Ie oi:

> oO:

. 43.
. 44,
. 40.

. 46.

pace
. 48.

49.

aeolle

PuatTE V.

General view of superficial blood vessels.
Slightly enlarged.

General view of arterial system. Slightly
enlarged.

Venous System and especially circulation in
Renal Organ. x 1.

Veins on external surface of Oviduct, ete.
ile

Prats VI.

Transverse section of Ctenidial axis. x 40.

Transverse section of Mucous gland. x 40.

Nervous system. x l.

Anterior Nerve centres, view from inside of
nerve collar. x 6.

Pedal Gangha in male. x 6.

Two leaflets of Osphradium. x 10.

Transverse section of osphradial leaflets.
nO: !

Cells from osphradium (maceration prepara-

tion). x 400.

Prats. VIL.

Transverse section through eye. x 140.

Cells from retina (maceration preparation).
x 400.

Outer wall of renal organ with renal filaments
seen from the lumen. «x l.

Nephridial Gland seen from inner surface
(facing lumen of renal organ). x 1.

Outer wall of renal organ after removal of
renal filaments, showing secondary lamellae.
xy as

Fig.

Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.

Fig.
Fig.
Fig.

Fig.
Fig.

ala.

52.
53.
54.
59.

56.

57a.
57b.
57e.

58.

09.

60.
61.

62.

63.

64.

65.
66.

BUCCINUM. 367

Diagram showing how blood passes to renal
filaments by vessels crossing the lumen of
the renal organ.

T.S. Renal filaments. x 200.

T.S. Secondary lamella. x 200.

Male gonad and ducts with penis. x 1.

Female gonoducts. x 1.

Puate VIII.

Section of tubule of testis showing formation
of spermatozoa. x l.

Spermatid.

‘* Hair-like ’’? spermatozoon

“ Worm-like ’’ spermatozoon after Retzius.

‘Transverse section of ovary. Very slightly

magnified.

Transverse section of vaginal part of oviduct.
Shghtly magnified.

Transverse section of ovary. x 150.

Under surface of egg capsule showing pore of
escape. x l.

Early stage in development of larva after
cannibalism (after Koren and Daniellsen).

Very early larval stage, otocysts present.
(after K. and D). |

Late veliger larva (original). x 50.

Egg capsules. Slightly reduced.

Shells of young Buccinum at time of departure
from egg capsules. x 10.

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