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TRANSACTIONS

LIVERPOOL BIOLOGICAL SOCIETY.

WOlG, 2SuUe

SESSION 1898-99.

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LIVERPOOL :
Prinrep sy T. Doss & Co., 229, BRowNLOW HILL.

1899.

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CONTENTS.

I. PROCEEDINGS.

Office-bearers and Council, 1898-99 : < oe
Report of the Council . : 7 val.
Summary of Proceedings at the Meetiies : Xi.
Laws of the Society ; Seve
List of Members . | x,
Librarian’s Report (with list of Edi cons ie TT) Xone

Treasurer's Balance Sheet . Preis Ss 2 LXSORIVE

II. TRANSACTIONS.

Presidential Address — ‘“‘The Relation between
Structure and Function, as Examined in the
meme by Prof. C. S. SHERRINGTON, M.A.,
Meer nS: ; ; i

Twelfth Annual Report of the Liverpool Marine
Biology Committee and the Biological Station
at Port Erin. By Prof. W. A. Herpman,

bse, ERS. . f 21
List of the Araneida of Port iin aaa Deen
By A. R. Jackson, B.Sc. (Vict.) : 45 266

Report on the Investigations carried on in 1898 in
connection with the Lancashire Sea-Fisheries
Laboratory at University College, Liverpool,
and the Sea-Fish Hatchery at Piel, near Barrow.

By Prof. W. A. Herpman, F.R.S., ANDREW
Scorr and JAMES JOHNSTONE , . 69

Iv. LIVERPOOL BIOLOGICAL SOCIETY.

Note on Mid-Winter Surface and Deep Tow-Nettings

in the Irish Sea. By Isaac C. THompson,
F.L.S.

Some Speculations on the Derivation of our British
Coleoptera. By W. HE. SHarp .

On Secondary Thickening in the Aerial Roots of
Hedera hex. By Prof. R. J. HARVEY GIBson,
NMeA, ELS.

Addendum to Report on a small Collection of
Antarctic Plankton, &c. By Isaac C. THompson,
ELS.

156

163

185

188

PROCEEDINGS

OF THE

LIVERPOOL BIOLOGICAL SOCIETY.

_e
i

OFFICE-BEARERS AND COUNCIL.

Ex-Presidents :

1886—87 Pror. W. MITCHELL BANKS, M.D., F.B.C.S.
1887—88 J. J. DRYSDALE, M.D.
1888—89 Pror. W. A. HERDMAN. D.S8c., F.B.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, O.M.Z.S., A.L.S.

189293 ALFRED 0. WALKER, J.P., F.LS.
1893—94 JOHN NEWTON, M.B.C.S.

1894 —95 Pror. F. GOTCH, M.A., F.B.S.

1895--96 Pror. R. J. HARVEY GIBSON, M.A.
189697 HENRY O. FORBES, LL.D., F.Z.8.
1897—98 ISAAC C. THOMPSON, F.L.S., F.R.M.S.

SESSION XIII, 1898-99.

President :
Pror. C. S. SHERRINGTON, M.D., F.R.S.
Pice-Presidents :
Pror. W. A. HERDMAN, D.Sc., F.R.S.
ISAAC C. THOMPSON, F.L.S., F.R.M.S.

How. Creasnrer :
T. C. RYLEY.
How. Hibrarian:
JAMES JOHNSTONE.

Hon. Secretary:
JOSEPH A. CLUBB, M.Sce., (Vicr.).

Council :
H. C. BEASLEY. | G. H. MORTON, F.G.S.
Ws J. HALLS. JOHN NEWTON, M.R.CS.
JOSEPH LOMAS, F.G.S. Pror. PATERSON, M.D.
Cuas. E. JONES, B.Sc. ©. RICKETTS, M.D.
Rey. L. de B. KLEIN, D.Sc. W. E. SHARP.
Rey. T. S. LEA, M.A. A. O. WALKER, F.L.S.

REPORT of the COUNCIL.

Durine the Session 1898-99 there have been seven
ordinary meetings and two field meetings of the Society.
The latter were held at Hilbre Island, jointly with the
Liverpool Geological Society and at Delamere Forest,
Cheshire, respectively. In response to an invitation from
the Owens College Biological Society, several of our mem-
bers attended one of the meetings at Manchester to hear
an illustrated address by Dr. Arthur Willey, entitled ‘‘ In
search of Nautilus,’ which embodied a description of his
recent travels in the Pacific.

The communications made to the Society have been
representative of almost all branches of Biology and the
exhibition of microscopic preparations and other objects
of interest has been well maintained at the meetings.

On the occasion of a lecture entitled ‘‘ Malaria and
Mosquitos,” given before the Society by Major Ross,
I.M.58., lecturer on Tropical Diseases at University College,
your Council adopted the plan of issuing special invitations
for the meeting, and the large and influential audience
which assembled testified both to the interest and the
value of the new departure.

The Library continues to make satisfactory progress,
and additional important exchanges have been arranged
during the year. The Hon. Librarian has in contemplation
the preparation of a detailed catalogue which will be of
ereat use to members. |

The Treasurer’s statement and balance sheet are
appended.

REPORT OF COUNCIL. 1X

No alterations have been made in the Laws of the
Society during the past session.
The members at present on the roll are as follows :—

Honorary Members ......... 9
Ordinary Members ......... 57
Student Members............ 16

SUMMARY of PROCEEDINGS at the MEETINGS.

The first meeting of the thirteenth session was held at
University College on Friday, October 14th, 1898.

1. The following exhibits were on view in the Zoological
Laboratory from 7 to 7-80.

A series of photographs, illustrating animal and
plant life between tide marks, was shown by Prof.
Herdman. 3

A number of mounted preparations from the
Fisheries Museum, by Mr. J. Johnstone.

The President-elect (Prof. C. 58. Sherrington, MIDE
F.R.S.) took the chair at 7-80, in the Zoology Theatre.

2. The Report of the Council on the Session 1897-98 (see
‘‘ Proceedings,” Vol. XII., p. vil.) was submitted
and adopted.

3. The Treasurer’s Balance Sheet for the Session 1897-98
(see ‘‘ Proceedings,” Vol. XII., p. xxxi.) was sub-
mitted and approved.

4. The Librarian’s Report (see ‘‘ Proceedings,” Vol. XI1.,
p- XXV.) was submitted and approved.

5. The following Office-bearers and Council for the ensuing
Session were elected :— Vice-Presidents, Prof.
Herdman, D.Sc., F.R.S., and I. C. Dhompeom
F.L.S., F.R.M.S; Hon. Treasurer, 1. G2 aigies
Hon. Librarian, James Johnstone; Hon. Secretary,
Joseph A. Clubb, M.Sc.; Council, H. C. Beasley,
W. J. Halls, Chas. H. Jones, B.Sc., Rey. diese
Beaumont Klem, D.Sc. Rev. TV. S. Heasyieas

SUMMARY OF PROCEEDINGS AT MEETINGS. xi

Joseph Lomas, F.G.8., G. H. Morton, F.G.S.,
John Newton, M.R.C.S., Prof. Paterson, M.D.,
Mii. Coo., ©. Rickets, M.D., W. H. Sharp, A. O.
Walker, F.L.S.

6, Prof. Herdman submitted the Twelfth Annual Report
on the work of the Liverpool Marine Biology
Committee and the Port Erin Biological Station
(see ‘* Transactions,” p. 21).

7. Mr. A. R. Jackson, B.Sc., gave a short note on his

work on the Spiders found in the neighbourhood of
Port Erin (see ‘‘ Transactions,” p. 66).

The second meeting of the thirteenth session was held
at University College on Friday, November 11th, 1898.
The President in the chair.

1. Prof. Sherrington, M.D., F.R.8., delivered the Presi-
dential address, entitled ‘‘ Adaptation to Function
as exemplified by the Structure of the Arm”’ (see
“Transactions,” p. 1). A vote of thanks was pro-
posed by Dr. Wiglesworth, seconded by Prof.
Boyce and carried with acclamation.

The third meeting of the thirteenth session was held at
University College on Friday, December 9th, 1898, and
took the form of a joint meeting with the Liverpool
Geological Society and the Historic Society of Lancashire
and Cheshire. The President in the chair.

1. In the Zoological Museum of the College Prof. Herdman

exhibited a series of Anthropological specimens and

Prof. Paterson described a series of exhibits,

- prepared by Dr. Lovegrove and himself, illustrating

peculiarities in the vertebral column of the human
foetus.

Xll. PROCEEDINGS LIVERPOOL BIOLOGICAL SOCIETY.

2. Mr. J. Lomas, F.G.S., delivered a lecture on ‘‘ Recent
Finds of Flints in Cheshire and North Wales, and
their bearing on the Antiquity of Man.’ A number
of the specimens found were exhibited and the
opinions of eminent experts were quoted to show
that these flints owed their shape to human rather
than natural agencies. The lecture was also illus-
trated by photographs and lantern slides. An
animated discussion followed.

The fourth meeting of the thirteenth session was held
at University College on Friday, January 13th, 1899.
Prof. Herdman (Vice-President) in the chair.

1. In the Zoological Laboratory Prof. Herdman exhibited
a series of microscopical preparations of the radulee
of molluscs, and Dr. Wiglesworth exhibited and
made some remarks upon the eggs of the Roseate
Tern (Sterna dougall).

2. A paper on ‘‘ Aerial Roots of the Ivy,” by Prof.
Harvey Gibson, was communicated by Prof,
Herdman (see ‘‘ Transactions,” p. 185).

3. Mr. W. E. Sharp communicated a paper on ‘‘ Some
Speculations on the Derivation of our British
Coleoptera’”’ (see ‘‘ Transactions,” p. 168).

The fifth meeting of the thirteenth session was held at
University College on Friday, February 10th, 1899. The
President in the Chair.

1. From 7 to 7-80 in the Physiological Laboratory a series
of miscellaneous exhibits were on view, and Dr.
Grunbaum described an apparatus for measuring
the vitiation of the air in a room.

SUMMARY OF PROCEEDINGS AT MEETINGS. Xlil.

2. Prof. Paul gave a lecture entitled ‘‘ Contributions to
the Histological Study of Dentine.’’ With the aid
of a series of lantern slides produced from micro-
photographs the development of dentine was graphi-
cally traced. Many of the slides from which the
illustrations were prepared were on view under
microscopes, and were inspected with great interest
at the close of the lecture. |

The sixth meeting of the thirteenth session was held at
University College on Friday, March 17th, 1899. The
President in the chair.

1. Mr. J. Johnstone exhibited with remarks a series of
microscopic preparations illustrating Spawn for-
mation in the Marine Mussel.

2. The Annual Report of the Sea-Fisheries Laboratory
for 1898 by Prof. Herdman, A. Scott, and J.
Johnstone was submitted (see ‘‘ Transactions,”’
eawg).

3. Mr. I. C. Thompson submitted some notes comparing
the results of Deep-Sea and surface Tow-Nettings
in the Irish Sea (see ‘‘ Transactions,” p. 156).

4. Prof. Paterson gave a lecture on ‘‘ Perforations in the
Parietal Bones, including an account of a micro-
cephalic cranium from the Liverpool Museum.” An
interesting series of photographs were shown of
the skulls with such perforations in illustration of
the lecture.

The seventh meeting of the thirteenth session was held
at University College on Friday, May 12th, 1899. The
President in the chair,

XIV. SUMMARY OF PROCEEDINGS AT MEETINGS.

1. A series of miscellaneous exhibits were on view previous |
to the meeting.

2. Major Ross, I.M.S., gave a lecture on ‘‘ Malaria and
Mosquitos,’ in which he showed the important
part played by mosquitos in the spread of malaria.
A beautiful series of microscopic sections illustrated
the lecture, which was listened to with great interest

by a large audience.

The eighth meeting of the thirteenth session was held
as a Field Meeting, jointly with the Liverpool Geological
Society, on Saturday, June 10th, 1899, at Hilbre Island
and West Kirby.

eee

The ninth meeting was held, also as a Field Meeting,
at Delamere Forest, Cheshire, on Saturday, June 17th.
After tea a short business meeting was held. On the
motion of Prof. Sherrington, seconded by Mr. Ryley, Dr.
Wiglesworth was unanimously elected President for next
session. On the motion of Prof. Herdman, seconded by
Mr. Beasley, a vote of thanks was passed, with acclama-
tion, to Prof. Sherrington the retiring President. —

LAWS of the LIVERPOOL BIOLOGICAL
SOCIETY.

I.—The name of the Society shall be the ‘‘ LIvERPooL
BionLocicaL Society,” and its object the advancement of
Biological Science.
Ii.—The Ordinary Meetings of the Society shall be held

at University College, at Seven o’clock, during the six
Winter months, on the second Friday evening in every
month, or at such other place or time as the Council may
appoint.

I11.—The business of the Society shall be conducted by
a President, two Vice-Presidents, a Treasurer, a Secretary,
a Librarian, and twelve other Members, who shall form a
Council; four to constitute a quorum.

IV.—The President, Vice-Presidents, Treasurer, Secre-
tary, Librarian, and Council shall be elected annually, by
ballot, in the manner hereinafter mentioned.

V.—The President shall be elected by the Council
(subject to the approval of the Society) at the last Meeting
of the Session, and take office at the ensuing Annual
Meeting.

VI.—The mode of election of the Vice-Presidents,
Treasurer, Secretary, Librarian, and Council shall be in the
form and manner following :—It shall be the duty of the
retiring Council at their final meeting to suggest the names
of Members to fill the offices of Vice-Presidents, Treasurer,
Secretary, Librarian, and of four Members who were not

XVl. LIVERPOOL BIOLOGICAL SOCIETY.

on the last Council to be on the Council for the ensuing
session, and formally to submit to the Society, for election
at the Annual Meeting, the names so suggested. The
Secretary shall make out and send to each Member of the
Society, with the circular convening the Annual Meeting,
a printed list of the retiring Council, stating the date of
the election of each Member, and the number of his atten-
dances at the Council Meetings during the past session;
and another containing the names of the Members sug-
gested for election, by which lists, and no others, the votes
shall be taken. It shall, however, be open to any Member
to substitute any other names in place of those upon the
lists, sufficient space being left for that purpose. Should
any list when delivered to the President contain other
than the proper number of names, that list and the votes
thereby given shall be absolutely void. Every list must
be handed in personally by the Member at the time of
voting. Vacancies occurring otherwise than by regular
annual retirement shall be filled by the Council.

VII.—Every Candidate for Membership shall be pro-

posed by three or more Members, one of the proposers
- from personal knowledge. The nomination shall be read
from the Chair at any Ordinary Meeting, and the Candi-
date therein recommended shall be balloted for at the
succeeding Ordinary Meeting. Ten black balls shall
exclude. .

VIII.—When a person has been elected a Member, the
Secretary shall inform him thereof, by letter, and shall at
the same time forward him a copy of the Laws of the
Society.

IX.—Every person so elected shall within one calendar
month after the date of such election pay an Entrance Fee
of Half a Guinea and an Annual Subscription of One

LAWS. XvVii.

Guinea (except in the case of Student Members) ; but the
Council shall have the power, in exceptional cases, of
extending the period for such payment. No Entrance
Fee shall be paid on re-election by any Member who has
paid such fee.

X.—The Subscription (except in the case of Student
Members) shall be One Guinea per annum, payable in
advance, on the day of the Annual Meeting in October.

XI.—Members may compound for their Annual Sub-
scription by a single payment of Ten Guineas.

XII.—There shall also be a class of Student Members,
paying an Entrance Fee of Two Shillings and Sixpence,
and a Subscription of Five Shillings per annum.

XIII.—All nominations of Student Members shall be
passed by the Council previous to nomination at an Ordin-
ary Meeting. When elected, Student Members shall be
entitled to all privileges of Ordinary Members, except
that they shall not receive the publications of the Society
nor vote at the Meetings, nor serve on the Council.

XIV.—Resignation of Membership shall be signified in
writing to the Secretary, but the Member so resigning shall
be liable for the payment of his Annual Subscription, and
all arrears up to date of his resignation.

XV.—The Annual Meeting shall be held on the second
Friday in October, or such other convenient day in the
month as the Council may appoint, when a Report of the
Council on the affairs of the Society, and a Balance Sheet
duly signed by the Auditors previously appointed by the
Council, shall be read.

XVI.—Any person (not resident within ten miles of
Liverpool) eminent in Biological Science, or who may have
rendered valuable services to the Society, shall be eligible

XVill. LIVERPOOL BIOLOGICAL SOCIETY.

as an Honorary Member; but the number of such Members
shall not exceed fifteen at any one time.

XVII.—Captains of vessels and others contributing
objects of interest shall be admissible as Associates for a

period of three years, subject to re-election at the end of
that time.

XVIII.—Such Honorary Members and Associates shall
be nominated by the Council, elected by a majority at an
Ordinary Meeting, and have the privilege of attending and
taking part in the Meetings of the Society, but not voting.

XIX.—Should there appear cause in the opinion of the
Council for the expulsion from the Society of any Member,
a Special General Meeting of the Society shall be called
by the Council for that purpose; and if two-thirds of those
voting agree that such Member be expelled, the Chairman
shall declare this decision, and the name of such Member
shall be erased from the books.

XX.—Every Member shall have the privilege of intro-
ducing one visitor at each Ordinary Meeting. The same

person shall not be admissible more than twice during the
same session.

XXI.—Notices of all Ordinary or Special Meetings shall
be issued to each Member by the Secretary, at least three
days before such Meeting.

XXII.—The President, Council, or any ten Members
can convene a Special General Meeting, to be called within
fourteen days, by giving notice 1n writing to the Secretary,
and stating the object of the desired Meeting. The circular
convening the Meeting must state the purpose thereof.

XXIII.—Votes in all elections shall be taken by ballot,

and in other cases by show of hands, unless a ballot be
first demanded,

LAWS. X1X.

XXIV.—No alteration shall be made in these Laws,
except at an Annual Meeting, or a Special Meeting called
for that purpose; and notice in writing of any proposed
alteration shall be given to the Council, and read at the
Ordinary Meeting, at least a month previous to the meet-
ing at which such alteration is to be considered, and the
proposed alteration shall also be printed in the circular
convening such meeting; but the Council shall have the
power of enacting such Bye-Laws as may be deemed neces-
sary, which Bye-Laws shall have the full power of Laws
until the ensuing Annual Meeting, or a Special Meeting
convened for their consideration.

BYE-LAWS.

1. Student Members of the Society may be admitted as
Ordinary Members without re-election upon payment of
the Ordinary Member’s Subscription; and they shall be
exempt from the Ordinary Member’s entrance fee.

2. University College Students may be admitted as
Student Members of the Society for the period of their
college residence, on the single payment of a fee of Five
Shillings and an entrance fee of Two Shillings and

Sixpence.

LIST of MEMBERS of the LIVERPOOL

ELECTED.

1898
1886
1886
1888
1894
1889
1886
1886
1897
1894

1891
1886

1886
1898

1886
1896

BIOLOGICAL SOCIETY.
SHSSION 1898-99.

A. ORDINARY MEMBERS.
(Life Members are marked with an asterisk.)

Armour, Dr. T. R. W., University College,
Liverpool

Banks, Prof. W- Mitchell, M.D., F.B.C.S., 28;
Rodney-street

Barron, Prof. Alexander, M.B., M.R.C.S., 34,
Rodney-street

Beasley, Henry C., Prince Alfred-road, Wavertree

Boyce, Prof. University College, Liverpool

Brown, Prof. J. Campbell, 8, Abercromby-square

Caton, R., M.D, F.R.C.P., Lea Hall, Gateacre

Clubb, J. A., M.Sc., Hon. SECRETARY, Free
Public Museums, Liverpool

Dutton, Dr. J. Everett, 502, New Chester-road,
Rock Ferry

Forbes, H. O., U.D., F.Z.S., Free ube
Museums, Liverpool

Garstang, W., M.A., Lincoln College, Oxford

Glynn, Prof. T. R., M.D., F.R.C.P., 62, Rodney-
street

Gibson, Prof. R. J. Harvey, M.A., F.U.8., Univer-
sity College

Grunbaum, Dr. A. §., 67, Rodney Street

Halls, W. J., 35, Lord-street

Haydon, W. H., 24, Upper Parliament-street

1886

1893

1891
1894

1897
1898

1886

1894

1895

1894
1896

1886

1888
1886
1888
1894

1894
1892
1896
1886
1897
1890
1895

LIST OF MEMBERS. XX1,

iterdman, Prof. W. A., D.Sc., F.R.8., Vice
PRESIDENT, University College

Herdman, Mrs., B.Sc., Croxteth Lodge, Ullet-road,
Liverpool

Hicks, J. Sibley, M.D-, 2, Erskine-street

iuekson, Prof. 5S: J., F.R.S., Owens College,
Manchester

Holt, Alfred, Crofton, Aigburth

Johnstone, James, Hon. LIBRARIAN, Fisheries
Laboratory, University College, Liverpool

Jones, Charles W., Field House, Prince Alfred-
road, Wavertree

Jones, Charles Elpie, B.Sc., Prenton-road, W..,
Birkenhead

Klein, Rev. L.. de Beaumont, D.Sc., F.L.5., 6,
Devonshire-road ;

ea, Rev. T. S., St. Ambrose Vicarage, Widnes

-Liaverock, W. 8., M.A., B.Sc., Free Museums,

Liverpool
Lomas, J., Assoc. N.8.8S., F.G.S., 16, Mellor-road,
Birkenhead
Melly, W. R., Ph.D., 90, Chatham-street
Morton, G. H., F'.G.S., 209, Kdge-lane, EH.
Newton, John, M.R.C.S., 44, Rodney-street
Paterson, Prof., M.D., M.R.C.8., University Col-
lege, Liverpool
Paul, Prof. F. T., Rodney-street, Liverpool
Phillips, E., L.D.8., M.R.C.S., 38, Rodney-street
Picton, W. H., 2, College-road, Gt. Crosby

*Poole, Sir James, J.P., Abercromby-square

Quayle, Alfred, 7, Scarisbrick New-road, S’port

*Rathbone, Miss May, Backwood, Neston

Ricketts, C., M.D., 11, Hamilton-square, B’head

XXil1.
1887

1897
1887

1894
1895

1891
1886

1895
1898
1893
1886

1889
1888
1886
SO7
1891
1896

1896

LIVERPOOL BIOLOGICAL SOCIETY.

Robertson, Helenus R., Springhill, Church-road,
Wavertree

Robinson, H. C., Holmfield, Aigburth

Ryley, Thomas C., Hon. TREASURER, 10, Waver-
ley-road :

Scott, Andrew, Piel, Barrow-in-Furness

Sherrington, Prof., M.D., FR), (earsmoamma
University College, Liverpool

Sharp, W. E., The Woodlands, Ledsham

Smith, Andrew T., Jun., 5, Hargreaves-rd., Sefton
Park

Smith, J., F.L.8., Rose Villa, Lachford, Warrington

Suffield, Miss, University College, Liverpool

Tate, Francis, F.C.S., 9, Hackins Hey, Liverpool

Thompson, Isaac C., F.L:S., ER Mess yen
PRESIDENT, 53, Croxteth-road

Thornely, Miss L. R., Baychiff, Woolton Hill

Toll, J. M., 49, Newsham-drive, Liverpool

Walker, Alfred O., J.P., F.L.8., Colwyn Bay

Warrington, Dr. W. B., 80, Rodney-street

Wiglesworth, J.. M.D., County Asylum, Rainhill

Woods, Joseph A., L.D.S. Eng., 76, Mount-
pleasant, Liverpool

Wilmer, Miss J. H., 20, Lorne-rd., Orton, B’head

B. StTuDENT MEMBERS.

Armstrong, Miss A., 26, Trinity-road, Bootle
Bennette, Jiorace W. P., Gothic Lodge, Park-road, 8.,

Birkenhead

Carstairs, Miss, Lily-road, Fairfield

Crompton, Miss C. A., University College, Liverpool
Dickinson, T., 3, Clark-street, Princes Park |
Drinkwater, E. H., Rydal Mount, Marlboro’-road, Tue-

brook

LIST OF MEMBERS. XX1ll.

Hlder, D., 49, Richmond Park, Liverpool

Gill, E. 8. H., Shaftsbury House, Formby

Hannah, J. H. W., 55, Avondale-road, Sefton Park
Harrison, Oulton, Denehurst, Victoria Park, Wavertree
Henderson, W. 8., B.Sc., Beech-hill, Fairfield

Knoit, Henry, 46, Underley-street, Liverpool

Lloyd, J. T., 48, Ullet-road, Sefton Park

Mann, J. C., University College, Liverpool

Mawby, W., Clumber, Prenton-road, EH, Birkenhead
Woolfenden, H. F., 6, Grosvenor-road, Birkdale

C. HonorRARY MEMBERS.

H.S.H. Albert I., Prince of Monaco, 25, Faubourg St.
Honore, Paris

Bornet, Dr. Edouard, Quai de la Tournelle 27, Paris

Claus, Prof. Carl, University, Vienna

Fritsch, Prof. Anton, Museum, Prague, Bohemia

Giard, Prof. Alfred, Sorbonne, Paris

Haeckel, Prof. Dr. EK., University, Jena

Hanitsch, R., Ph.D., Raffles Museum, Singapore

Leicester, Alfred, Buckhurst Farm, nr. Edenbridge, Kent

Solms-Laubach, Prof- Dr., Botan. Instit., Strassburg

REPORT of the LIBRARIAN.

——

DurineG the last session of the Society, retrospective ex-
changes, including most of the former volumes of the
Transactions, were arranged with the Kansas University
(U.S.A.) and the Illinois (U.8.A.) State Laboratory of
Natural History.

Attention was directed in last year’s Report to the ae
proportion of journals and other publications in the
Society's Library which still remain unbound, and which
has been largely increased by last year’s additions. Until
this is done the arrangement and cataloguing of the
Librarv is not convenient nor indeed practicable.

Lists are given below of the publications which have
been added, by exchange and otherwise, to the Library
since the end of the Twelfth Session, and of the Societies
and Institutions with whom publications are exchanged.
1. Batavia, Naturkundig Tijdschrift v. Nederlandsch-Indié. Deel LVII.,

Tiende Serie, Deel I. 1898.
2. Boston, (U.S.A.) Proceedings of the Boston Society of Natural History,
Vol. 28, pp. 117—800, December, 1897—July, 1898.
3. Bonn, Sitzungsberichte d. Niederrheinschen Ges. f, Natur-und Heilkunde.
Erste und Zweite Halfte. 1898.
4. Bonn, Verhandl. d. Naturhist. Vereins d. Preussisch. Rheinlande.
Funf. Jahr. Erste u. Zweite Halfte. 1898.
5. Bologna, Memorie d. R. Accademia d. Scienze dell’ Instituto di Bologna.
Serie V.—T. VI. Sezione. di Scienze Naturali. Sezione di
Medicino e Chirurgia. 1896-97.
6. Bordeaux, Procés-verbaux de la Société Linnéenne de Bordeaux. Vol.
LII. 1897.

. Bergen, Bergens Museums Aarbog for 1898. 1899.

. Bergen, An account of the Crustacea of Norway. Vol. II. Isopaaee pts.
9—10, Munnopsidae. (concluded) Ligiidae, Trichoniscides Oniscidae
(part). G. O. Sars. 1898.

CO

LIBRARIAN'S REPORT. XXV.

9. Berlin, Sitzungsberichte der koniglich Preussischen Akademie der
Wissenschaften. I.—LIV., 1898.

Bristol, Publications of the British Association, Bristol Meeting, 1898.
(Presented by Prof. Herdman.)

10. The President’s Address.

alse Address to the Zoological Section by Prof. W. F. R. Weldon, F.R.S.

12. Address to the Botanical Section by Prof. F. O. Bower, F.R.S.

13. Life Conditions of the Oyster, 8rd Report of the Committee, by Prof.

Herdman, Prof. Boyce, and Dr. Kohn.
14. Preliminary Note on the Migrations of the Mackerel. W.
Garstang, M.A.

15. Buenos Aires, Comunicaciones del Museo Nacional de Buenos Aires.
Homovt, No. 1, 2. 1898.

16. Brussels, Annales du Musée du Congo. Serie I.—Botanique ; Tome. I.
Fase. 1 and 2. Illustrations de la Flora du Congo. Em. de
Wildeman et Th. Surand. 1898.

Serie II., Zoologie, Tome I., Fasc. 2. Materiaux pour la Faune du
Congo. Poissons Nouveaux. G. A. Boulenger. 1898. (Presented
by the Secretary of State, Congo Free State. )

17. Cambridge, Mass., U.S.A., Bulletin of the Museum of Comparative
Zoology at Harvard College. Vol. XXXII., Nos. 1—9; Vol. XXVIII.,
Nos. 4—5. (Geological Series Vol. III.).

18. Cambridge, Mass., U.S.A., Annual Report of the Curator of the Museum,
Comparative Zoology at Harvard. 1897—98.

19. Charlottenburg, Zeitschrift fiir Fischeri u. deren Hilfswissenschaften mit
einschluss v. Fischwasser-Hygiene, Fischerei u. Wasserrecht. Heft
1—5, 1898.

20. Charlottenburg, Mitglieder- Liste des Deutschen Fischerei - Vereins.
October 1, 1898.

21. Copenhagen, Beretning fra. Kommissionen f. Vidensk. Undersogelse af
de Danske Farvande. Andet. Bind. Forste. v. And. Heft.
1897-99.

22. Copenhagen, Oversigt over det Kongelige Danske Videnskabernes
Selskabs Forhandliger. Nos. 1--6, 1898-99.

23. Copenhagen, Videnskabelige Meddelelser fra den Naturhisoriske Forening
i Kjobenhavn f. Aaret.. 1898.

24. Christiania, Forhandlinger i Videnskabs-Selskabet i Christiania. Aar
1897-98.

25. Dublin, Scientific Proc. Roy. Dublin Soc. Vol. VIII. (N.S.). Novem-
ber, 1898, pt. 6.

26. Dublin, The Scientific Trans. Roy. Dublin Soc. Vol. V. (Ser. II.)—
MaAVe—KVI.; Vol. VI., I.

?

XXVI1. LIVERPOOL BIOLOGICAL SOCIETY.

27.

28.

29.

30.

3l.

32.

33.

o4,

35.

36.

37.

38.

39.

40.

41.

42.
43.

44,

45.

46.

Edinburgh, Proceedings of the Royal Society of Edinburgh. Vol. XXL.,
Sessions 1895-96, 1866-97.

Edinburgh, Sixteenth An. Report of the Fishery Board for Scotland,
1897. Pt. 3. Scientific Investigations.

Edinburgh, Report of Investigations on the Life History of the Salmon.

Fishery Bd. for Scotland.

Freiburg, Berichte der Naturforschenden Gesellschaft. Bd. X. Hefte
1—3.

Frankfurt, Bericht d. Senckenbergischen Naturforschenden Gesellschaft.
June, 1897, to June, 1898.

Frankfurt, Katalog d. Reptilien-Sammlung im Museum d. Senckenberg
Naturf. Ges. i. Frankfurt am Main. Teil II. (Schlangen). Prof. ~
Dr. O. Boettger. May, 1898.

Glasgow, Transactions of the Natural History Society of Glasgow. Vol.
WeguCNES: ae lat-e2. el SO7-98:

Geneva, Mem. de la Soc. de Physique et d’ Historie Naturelle.
T. XXXIII. Premiere Partie. 1898.

Gottingen, Nachrichten v.d. Konig. Gesell. d. Wissenschaften. Math-
phys. Klasse. Heft 1, 4, 1898. Geschaftliche Mitt, 1898, Heft I.

Haarlem, Archives du Musee Teyler. Ser. II., Vol. VI., 1, 2, and 4,
1898.

Hannover, Mitth. d. Deutschen Seefischerei-Vereins. Bd. XIV., Nos.
2-12.

Hannover, Fischerei-Zeitung. Sept. 28, 1898.

Havana, Tuberculosis en la Habana. Dr. A. de Gordon y de Costa.
Indicaciones Terapeuticas de la Musica. Dr. A. de Gordon y de
Costa (9 and 10 presented by the Author).

Havana, La Legislacion del Seguro de Vita ante la Medicina Forsene.
Dr. Antonio de Gordon y de Acosta. (From the Author). 1898.
Helsingfors, Das Thierleben im Nurmijarvi-See. Eine faunische-

biologische Studie. Von K. E. Stenroos. 1898.

Jersey, The Journal of Marine Biology, Vol. II., No. 6. Dec. 1898.

Kiel u. Leipzig, Wissenschaftliche Meeresuntersuchungen. Komm. z.
Wissenschaftlichen Untersuchung d. Deutsch Meere i. Kiel u.d.
Biologischen Anstalt auf Helgoland. Neue Folge. 2B. III.
Abtheilung Kiel. 1898.

Lawrence, U.S.A., The Kansas University Quarterly. Vols. I. to VIII.
1892-99.

La Haye, Archives Néerlandaises des Sciences Exactes et Naturelles.
Serie II., Tome 2. Liv. 1—4, 1898-99.

La Plata, Revista del Museo de La Plata. Tomo. VIII., 1898.

47.

48.

49.

50.

51.

52.

53.

54.

55.

57.

58.

59

60.

LIBRARIAN’S REPORT. XXVil.

Leipzig, Berichte u.d. Verhandlungen d. Konig]. Sachs. Gesellsch. d.
Wissensch. Mathematisch-physische Classe. Bd. 50, 1898,
Naturwissenscht. Theil. 1899. Math.—Theil V. 1898.

Leipzig and Vienna, Kunsten-Formen der Natur. Erste Lieferung, von
Ernst Haeckel. (Presented by the Author.)

Limoges, Etudes sur les Fourmis les Guépes et les Abeilles. C. Janet.
Note 13, Sur le lasius mixtus, ’antennophorus ulhmanni.

Note 14, Rapports des animaux myromecophiles avec les fourmis.

London, Journal of the Royal Microscopical Society, parts 3 —6, 1898 ;
parts 1—2, 1899.

London, The Naturalist. Nos. 497—-508, 1898; Nos. 504-508, 1899.

London, On the Heart-Body and Celomic Fluid of certain Polycheta.
ieeeicton, BoA., reprinted from @.J.M.Se.— Vol. 41, part 2;
(N.S.). (Presented by the Author.)

London, On some Crustaceans from the South Pacific, part 2., Macrura
anomala. J,. A. Borradaile, M.A., F.Z.S., reprinted from Proc.
Zool. Soc., No., XXXI., 1898. June, 1898.

London, A Revision of the Pontoniidae. L. A. Borradaile, M.A., F.Z.S.
reprinted from An. and Mag. Nat. Hist. Ser. 7, Vol. II., Nov.,
1898. Nos. 3 and 4 presented by the Author.

London, Hippolyte fascigera, Grosse, and H. gracilis, Heller. A. O.
Walker. Annals and Mag. Nat. Hist. Ser. 7. Vol. III. 1899.
(Presented by the Author.)

London, Catalogue of Fishes in the British Museum (N.H.), 2nd edition,
Vol. I. The Perciform Fishes (part). G. A. Boulenger, F.R.S.
1895.

Liverpool, Proceedings of the Liverpool Geological Society. Session 39.
1897-98, part 2, Vol. VII. 1898.

Liverpool, On the Floor Deposits of the Irish Sea. W. A. Herdman,
F.R.S., and J. Lomas, A.R.C.S., reprinted from Proceedings,
Liverpool Geological Society, 1897-98. (Presented by the Authors.)

. Liverpool, Bulletin of the Liverpool Museums. Vol. I. No. 38—4.

October, 1898.

Liverpool, Buried Bones about Liverpool. The Elephant in Cheshire.
Transactions Liverpool Biological Society. XII. 1898. G. H.
Morton. (Presented by the Author.)

Madison, Wisconsin, U.S.A., Wisconsin Geological and Natural History
Survey. Bull. I. Economic Series I.

On the Foresty Conditions of Northern Wisconsin. Filibert Roth.
1898.
Bull. II. Scientific Series I.

XXVII1. LIVERPOOL BIOLOGICAL SOCIETY.

(oy)
bo

68.

69.

Madison—
On the Instincts and Habits of the Solitary Wasps. G. W. Peckham
and Elizabeth G. Peckham. 1898.

Madison, Wisconsin, U.S.A., Transactions of the Wisconsin Academy.
Vol. XI., 1896—97, 50 plates. 1898.

Manchester, Transactions and Annual Report, Manchester Microscopical
Society, 1897.

Meriden, Conn., U.S.A., Transactions of the Meriden Scientific Associa-
tion, 1897—98.

Melbourne, Proceedings of the Royal Society of Victoria. Vol. XI., (U.S.)
part 1. 1898.

Monaco, Résultats des Campagnes Scientifiques accomplies sur son Yacht
par Albert Iet Prince Souverain de Monaco. Fase. XII. Echinides
et Ophiures. (Golfe de Gascogne, Acores, Terre- Neuve, 1898.)
(Donation. )

Moscow, Bulletin de la Société Impériale des Naturalistes. Nos. 3, 4,
1897 ; Nos. 1—3, 1898.

Monte Video, Anales del Museo Nacional de Montevideo. Tomo. II.,
Fase. VIII., Tomo. III., Fase. VIIJ. and X., 1898. °

Munich, Allgemeine Fischerei-Zeitung. Nos. 9—23, 1898; Nos. 1-8,
1899.

Naples, Rendiconto dell. Accademia d. Scienze Fisiche e Matematiche.
Ser. 3. Vol. IV. (anno xxxvii.). Fasc. 3—12, 1898. Ser. 3a, Vol.
V. Fasc. 1— 3, 1899.

Nancy, Bull. de la Societe des Sciences dc Nancy. Ser. II.; T. XV. ;
Fase. XXXII., 30¢ an.—1897. 1898.

New York, Additional Characters of the Great Herbivorous Dinosaur
Camarasaurus. H. F. Osborn. (Reprinted from Bulletin of the
American Museum of Natural History. Vol. X., pp. 219—233),
June, 1898.

New York, A Complete Skeleton of Teleoceras Fossiger. Notes on the
Growth and Sexual Characters of this Species. H. F. Osborn. (Bull.
Am. Mus. Nat. Hist., pp. 51—59.) 1898. (35 and 36 presented by
the Author.)

New York, Zoology at Columbia, 1891-97, from Columbia Univ. Bull.
(Presented by H. F. Osborn.)

New York, Trituberculy, a review, H. F. Osborn. (Presented by the
Author.)

New York, Organic Selection. H. F. Osborn, from Science N.S. Vol.
VI., No. 146. (Presented by the Author.)

Paris, Bulletin du Museum d Histoire Naturelle. No. 7—8, 1897 , Nos.
1—6, 1898.

78.
a.
80.

81.

82.

83.

84.

96.

97,

LIBRARIAN’S REPORT. XX1X.

Paris, Mémoirés de la Société Zoologique de France. Tome X. 1897.

Paris, Bulletin de la Société Zoologique de France. Tome XXII. 1898.

Paris, Bull. Scientifique de la France et de la Belgique. T. XXVIII.,
Poamuanbien sos. LT. XXX). 1898.

Paris, Etudes sur les fourmis les guépes et, les abeilles. Note 15.
Appareils pour lVobservation des fourmis et des Animaux Myrmeéco-
philes. C. Janet. (Extrait des Memoires de la Soc. Zool. de
France.) 1897.

Paris, Notice sur les Travaux Scientifiques preséntés par M. C. Janet a
Vacademie d. Sciences au Concours de 1896 pour le prix thore.

Paris, Sur les rapports de ?Antennophorus uhlmanni Haller, avec le
Lasius mixtus Nylander. (Hxtrait des Com. Ren. hebd. des Seances
de l’Acad. des Sci. T. 124, p. 588, 1897.) (40, 41, 42, presented by
the Author.)

Paris, Sur l Ethologie du Campanularia caliculata. Hincks. (Stolon-
isation et allogonie.) A. Giard. (Extrait des Comp. Rend. d.
Seances de la Soc. de Biol. Jan., 1898.)

Philadelphia, Proceedings of the Academy of Natural Sciences, part 3,
1897 ; parts 1--3, 1898-99.

Porto, Annaes de Sciencias Naturaes. Anno—V—Vol. V., Nos. 1—8.

Rochester (U.S.A.), Journal of Applied Microscopy (specimen number).
Molt No. 11. 1898.

Santiago de Chili, Revista Chilena de Hijiene. Tome IV., Cu. I. 1898.

Santiago de Chili, Bolletin de Hijiene i Demografia. Afio I., Num. I.
1898.

Santiago de Chili, Sesiones del Consejo Superior de Hijiene Publica.
1897.

San Francisco, Proceedings of the California Academy of Science. Vol. I.
Zoology, Nos. 6—9. Geology, No. 4. Botany, Nos. 3—5. Mathe-
matics and Physics, Nos. 1—4.

Stavanger, Stavanger Museum. Aarsberetning for 1897.

Sydney, Records of the Australian Museum. Vol. III., No. 4. 1898.

Sydney Catalogue of the Australian Birds in the Australian Museum.
Parts I. and III. Accipitres and Striges. E, P. Ramsay and A. J
North. 1874-98.

Sydney, Australian Museum. Report of Trustees for the year 1897-
BON:

St. Louis (U.S.A..), Transactions of the Academy of St. Louis. Vol.
VII., Nos. 17—20. Vol. VIII., Nos. 1—7. 1897-98.

St. Petersburg, Bull. de l’Academie Imp des Sciences. V Serie T. VII.
Nos. 2—5, Oct., Dec., 1897, T. VILI., Nos, 1—4, Jan.—April,
1898, 1898,

‘XXX. LIVERPOOL BIOLOGICAL SOCIETY.

98. Stockholm, Bihang till Kong. Svenska Vetenskaps — Akad. handl.
Bd. 23. Afd. III.—IV., No. 7. 1898.

99. Torino, Bolletino dei Mus. di. Zool. ed. Anat. comp. Vol. XIII., N.
311—334. 1898.

100. Tokyo, Journal of the College of Science. Imperial University of
Japan. Vol. IX., pt. 3; Vol..X., pt. 35 Wolk ay spiel
XII., pts. 1—3. 1898-99.

101. Tokyo, Annotationes Zoologice Japonenses. Vol. II., pts. 1—4. 1898.

102. Toronto, Proceedings of the Canadian Institute. Nos, 4—5. Vol. I.,
pts. 4-5. No. 6, Vol. I., pt. 6. 1898.

103. Toronto, Trans. Canadian Institute. No. 10. Vol. I., part 2. 1898.

104. Tiflis, Bericht v. d. Kaukasische Museum y. d. Offentliche Bibliothek
in Tiflis. Fur die Jahre, 1897 v. 1898. 1898.

105. Tiflis, Mittheilungen des Kaukaisischen Museums. Bd. I. Lief. 1.

106. Upsala, Studien o Nagra Svenska vaxters groningsted och Forstarkn-
ingsstadium. Astrid Cleve. Akademisk Afhandling. 1898.

107. Upsala, Universitets Arsskrift. 1897. Medicin. I. Om vissa Organ-
extrakts Inverkau a det isolerade och ofverlefuande daggdjurshyartat.
K. Hedborn.

108. Upsala, Universitets Arsskert. 1898. Program. I. 1897.

109. Bidrag till en Lefnadsteckning ofver Carl von Linné, VII. Th. m.
Fries. 1898.

110. Upsala, Meddelanden fran Kongl. Landbruksstyrelsen. No.1. Ar.,
1898 (No. 43).

111. Undersokningar rorande, Oresunds Djurlif. Dr. E.Lounberg. 1898.

112. Upsala, Anatomiska Studier ofver Sydamerikanska Peperomier. Aka-
demisk Afhandling. E. Jaderholm. 1898.

113. Upsala, Bidrag till Kannedomen om de Svenska Fanerogama Osternas
Skottutveckling och ofverbintring. Akademisk Afhandling. J. A.
Z. Brundin. 1898.

114. Upsala, Nova Acta Regiae Societatis Scientiarum Upsaliensis. Ser. IV.
Vol. XVII.: Fase: IT. .1898.

115. Upsala, Koniklijk Zool. Genootschap ‘‘ Natura Artis Magistra.” 1838.
May 1, 1898. 1898.

116. Vienna, Verhandlungen d. Kaiserlich-Konigl. Zool. — Botanish.
Gesellsch. Jahrg., 1898. Bd. XLVIII. 1898. ;

117. Washington, Bulletin of the U.S. Nat. Museum. No. 47. The Fishes
of N. and Middle America. Pts.2and3. By D.S8. Jordan, Ph.D.,
and B. W. Ivermann, Ph.D. 1898.

118.. Washington, Proceedings of the U.S. Nat. Museum. Vol. XX.,
p-p. 775—901 and Index. Vol. XXI., Nos, 1140—1147, 1149—
1170.

LIBRARIANS REPORT. XXX1.

119. Washington, U.S.A., Report of the Commissioner of the U.S. Com. of
Fish and Fisheries. Part 22 for the year ending June 30, 1897.
Issued 1898.

120.. Washington, Bull. U.S. Fish Commission. Vol. XVII. 1897-1898.

121. Wellington, Trans. and Proc. of the New Zealand Institute, 1897.
Wiles (MIS N.S.) 1898.

122. Ziirich, Vierteljahrsschrift: d. Naturf. Ges. i Zurich. Jahrg. 48.
Hefte I.—IV. 1898-99.

List of Societies, etc., with which publications are
exchanged :—

Ams1eERDAM—Koninklijke Akadamie van Wettenschappen

Koninklijke Zoologisch Genootschap Natura Artis Magistra
BatrimorE—Johns Hopkins University
BaraviA—Koninklijke Natuurkundig Vereeniging in Ned. Indie.
Brrgen—Museum
Brrtin—Konigl. Akademie der Wissenschaften

Deutscher Fischerei-Vereins
BirMINGHAM—Philosophical Society
Bo.onga—-Accademia della Scienze
Boxnn—Naturhistorisclier verein des Preussichen Rheinlande und Westfalens
Borpnaux—Société Linnéenne
Boston—Society of Natural History
BrussEL.s —Academie Royal des Sciences, etc., de Belgique
Bunnos AiREsS—Museo Nacional
Museo de la Plata

CAMBRIDGE —Morphological Laboratories
CamBripGr, MAss.—Museum of Comparative Zoology of Harvard College
Cuaristi1AniA—Videnskabs-Selskabet
Dusiin—Royal Dublin Society
EpinBurGH—Koyal Society

Royal Physical Society

‘Royal College of Physicians

Fishery Board for Scotland
FRANKFURtT— Senckenbergische Naturforschende Gesellschaft
Freipurc-—Naturforschende Gesellschaft
GENEVE — Société de Physique et d’Histoire Naturelle
GiesstNn—Oberhessische Gesellschaft fiir Natur und Heilkunde
G.iascow— Natural Iistory Society
GorrinceN—Konigl. Gesel!schaft der Wissenschaften
Hauirax—Nova Scotian Institute of Natural Science

XXXil. LIVERPOOL BIOLOGICAL SOCIETY.

HARLEM—Musée Teyler
Société Hollandaise des Sciences
HELIGOLAND—KoOnigliche Biologische Anstalt
*Iuuinois, U.S.A.—Reports of the State Laboratory of Natural History.
KireL—Naturwissenschaftlichen vereins fur Schleswig—Holstein
Kommission fur der Unterschung der Deutschen meere

KsoBpenHAVN—Naturhistorike Forening

Danish Biological Station (C. G. John Petersen)

Kongelige Danske Videnskabernes Selskab
*Lawrence, U.S.A.—The Kansas University Quarterly.
Lrrps—Yorkshire Naturalists’ Union
Lrrezic—Konigl. Sachs. Gesellschaft der Wissenschaften
Littr—Revue Biologique du Nord de la France
Liverpoor—Geological Society

Bulletin of the Liverpool Museum
Lonpon—Royal Microscopical Society
British Museum (Natural History Department).
MANCHESTER—Microscopical Society
Owens College
MARSEILLES

Station Zoologique d’Endoume
Musée d’ Historie Naturelle
Massacuuserrs—Tufts College Library
MECKLENBURG
MeLBouRNE—Royal Society of Victoria
MonrevipEo—-Museo Nacional de Montevideo
Montprre._LiIer —Académie des Sciences et Lettres
Moscou—Société Impériale des Naturalistes
Nanoy—Société des Sciences
Napoti—Accademia delle Scienze Fisiche e Matematiche
New Brunswick—Natural History Society
Oporro—Annaes de Sciencias Naturaes
Parts—Museum d’Histoire Naturelle

Société Zoologique de France

Vereins der Freunde der Naturgeschichte

Bulletin Scientifique de la France et de la Belgique
PHILADELPHIA —Academy of Natural Sciences
PLtymourH—Marine Biological Association
Sr. Louis, Miss.—Academy of Sciences
Sr. Pererspurc—Académie Impériale des Sciences
San Franctsco--California Academy of Science
SANTIAGO—Société Scientifiq du Chili
STaVANGER-—Stavanger Museum

Those marked with * are the additions,

uae
—— a.

ee

Wie,

LIBRARIAN’S REPORT. XXXlll.

SrockHotmM—Académie Royale des Sciences
Sypnry—Australian Museum
Tox1o—Imperial University
Zoological Society of Tokyo
Tortno— Musei de Zoologia ed Anatomia Comparata della R. Universita
ToRon'1'0 —Canadian Institute
TriEsTE—Societa Adriatica de Scienze Naturali
UpsaLta—Upsala Universitiet.
Société Royale des Sciences
WASHINGTON—Smithsonian Institution
United States National Museum
United States Commission of Fish and Fisheries
WELLINGTON, N.Z.—New Zealand Institute
Wien—K. K. Naturhistorischen Hofmuseums
K. K. Zoologisch—Botanischen Gesellschaft
ZuricH—Zurcher Naturforschende Gesellschaft

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TRANSACTIONS

OF THE

RP OL BIOLOGICAL SOCIETY.

INAUGURAL ADDRESS
ON
THE RELATION BETWEEN STRUCTURE and
FUNCTION, as EXAMINED in the ARM.

By Pror. C. 8. SHerrineton, M.A., M.D., F.R.S., Presipent.
[Read November 11th, 1898].

In choosing the above for the title of the Address which,
as President for the year, I have the privilege to lay
before our Society, the term structure has had reference
in my mind not to physical or chemical structure so much
as to morphological. Morphology has as its object the
study of the form of living things, and comparative
anatomy it pursues as one of its best and most valuable
methods. Of biological studies, those in comparative
anatomy are amongst the oldest. The old masters, in
pursuing them, delighted to indulge in speculations con-
cerning the use of the structures they described. As the
various parts of the mechanisms whose form they
examined became known to them, they often had cause to
note the suitability of the instrument to its purpose in the
life of the creature. They frequently digressed from the
immediate object of their treatises to discourse upon the
evidences of design in creation of which their observa-
tions gave them proof. The Bridgewater Treatises were
founded in part to illustrate the beneficent design testified
to by the mechanism and vital endowments of the animal
body. One of the most famous of these well-known essays
was, and has remained, Sir Charles Bell’s treatise ‘‘On the
Hand,’”’ Its stately language and wealth of illustration

2 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

will long preserve it as a masterpiece of popular exposi-
tion of science. Yet every chapter seems far from us
as a bygone age; its pages are alive no longer. How
has this come about? The facts are true, and therefore
living as ever; the style is classical, and therefore never
dying. It is the point of view that has suffered change.
The question asked throughout is the question that it
was the fashion at that time for biologists, and especially
anatomists, to ask and to attempt to answer. It was of a
phase that was passed through by such students at the
period of the ‘‘encyclopcedists’”’ and of Rousseau, and
lingered for a generation longer. The question asked was
one beyond the limit of regions accessible by the means of
enquiry that obtains in natural philosophy. It is now
generally acknowledged that this kind of teleology lies
beyond the province of biology. We desire not to trespass .
across that limit. We are content to struggle with a
humbler problem. The question why? is not answered
by positive science, but only the question how ? and some-
times the question how much? The physiologist cannot
say why a muscle contracts, nor define “life.” To dog-
matise concerning the ‘‘why”’ of a bird’s flight, implies
the knowing the ‘‘ why”’ of the bird’s existence. We may
be able to see how things have happened, or how they
will happen; and it is a first step in the acquisition of
positive knowledge to know that the ratio ret is not the
‘‘yeason why.” ‘To confuse natural science with meta-
physics is a mischievous mistake for the enquirer, at
least for the present and for some centuries to come.

At first sight, function seems in many instances more
obviously related to morphological structure than is borne
out by a more searching examination of the two. Es-
pecially do writings upon mammalian anatomy furnish
numerous examples of the looseness of the logic that is

RELATION BETWEEN STRUCTURE AND FUNCTION. 3

considered sufficient to connect the two. The arm and |
hand offer a good field for an attempt to institute an
enquiry into the connection between the two. I will,
therefore, ask you to follow a brief sketch of the
morphological structure, and some remarks upon their
function; and that done, each can enquire of himself to
what measure the morphological structure has helped
toward the understanding of the eee of the perform-
ance of the function.

If you look at this outline of the human body you will
note that in the chest and trunk at least, there is quite
evidently a segmental arrangement of the body’s structure.
If you turn to this picture of the spinal cord lying in the
canal of the vertebrae, its composition out of a fore-and-aft
series of segments is still more obvious. The segments
or metameres of the body appear to possess one spinal
nerve apiece. If now we turn to this photograph of the
nerves passing from the spinal cord into the limb, we see
that a short series of these spinal segmental nerves all go
into it. This suggests that the limb is like the trunk
composed of metameres essentially arranged segmen-
tally, and built up asisthe rest of the body. But there is a
difficulty in deciding what part of the limb is constituted by
this or by that segment. 'The segmental nerves that issue
from the spinal cord into the limb commingle together
in a network or plexus. The brachial plexus, which is
before you in the photograph, is compounded of five,
sometimes of six spinal nerves. The nerve-trunks that
issue from it to the muscles and other structures of the
limb have obtained their nerve-fibres from various of the
spinal nerves, and each of them from several. To unravel
by gross anatomy, this intricate plexus is impossible.
Professor Krause, the anatomist of Berlin, has further
pointed out that not only was he unable to unravel

4 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the plexus, but. he found that each of the great nerve-
cords descending from the plexus into the limb 1s itself a
plexus, that defies dissection.

Histology (microscopic anatomy) and physiology can,
however, by reason of their finer methods overcome
the difficulty thus offered. The physiologist, John Muller,
discovered early in this century the ‘‘law of isolated
conduction ’”’ of nerve-fibres. If, therefore, a spinal nerve

be excited, e.g., by an electric current conveyed to it as it.

leaves the spinal cord before it enters the plexus, the fibres
that are contributed by it to the various nerves of the limb,
will be excited and no other fibres, however close its own
may run to them. :

The functions of the arm and hand are, however, so
multitudinously various that it is well to choose some
single, simple one, and to regard it only, letting it stand
as a concrete instance and subject for the physiological
part of the enquiry. Such an instance may well be chosen
in the simple movement I perform when holding my arm
out horizontally I flex my fingers, folding them into the
palm. ‘The question before us therefore is how is that
movement performed, and how does the morphological
structure of the part assist us toward explaining in what
-way the mechanism is actuated ?

. The flexion of the fingers is due to the contraction of
certain muscles situated on the ventral aspect of the forearm
and in the palm. In order to simplify our problem still
further, let us exclude the palmar muscles. The move-
ment can certainly be performed—as has been proved
experimentally in the monkey—without the contraction of
the palmar muscles, and this is a further justification for
simplifying our problem by their exclusion. The
muscles in question, in the forearm, are thrown into con-
traction by the action. of the nerves which pass to them

]
|
|
:
.

RELATION BETWEEN STRUCTURE AND FUNCTION. 5

from the spinal cord. These nerves are bundles of nerve-
fibres, while each one of them is a branch of a nerve-cell
lying within the spinal cord. The course of these fibres
gives important information as to the morphological
arrangement of the structure of the limb.

In this way can be discovered exactly what parts of the
musculature of the limb are innervated by each segmental
nerve. Another plan of investigation is based on the law
of degeneration. When an animal cell—for instance, one
of the large Amoebae obtained in the Red Sea—is divided
into several pieces, by tearing with a needle under a dis-
secting lens, each of the several pieces isolated draws
itself together, performs movements, and at first appears
to be capable of leading an independent existence. Con-
tinued observation, however, reveals the fact that the
isolated pieces after a few days die and disintegrate, with
the exception only of that portion in which the nucleus
remains; that lives on, and later repairs its injured bulk.
It is the same with nerve-cells, and probably with all
other animal cells. In the case of nerve-cells the
occurrence of degeneration in the portion of a cell which
has been isolated from the nucleus-containing portion of
the cell is, perhaps, the most valuable of all methods for
tracing the nerve-fibres to their parent nerve-cells.

The method of degeneration can be combined with that
of excitation by electricity. In a few days after nerve-
fibres have been severed from their parent nerve-cells, the
nerve-fibres, even before they exhibit visible signs of decay,
have lost their capacity to respond to stimuli and to con-
duct the waves of change which are called nervous impulses.
In this circumstance lies a means of escape from the em-
barrassing, for the investigator, accidents of ‘‘ escape of
current.”” The escape of the electric current from that
particular nerve to which it is intended. to solely apply, it

6 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

is difficult to avoid when these lie in the immediate neigh-
bourhood of other nerve trunks. By rendering these
latter inexcitable by reason of their degeneration, the
source of fallacy can be completely set aside.

In the above, great stress is obviously laid by implication
upon the distribution of the nerves as a key to the mor-
phological, especially to the segmental or metameric
architecture of the limb. Why are the nerve-fibres such
extremely important indices in this problem? The nerve-

cells in the mammal, at least, are peculiar in not evincing

reproductive activity after a very early period in the
individual’s hfe. They are early differentiated, and are like a
committee formed without power to add their numbers.
They are laid down when the metamerism of the relatively
simply-shaped young individual is more easily traced than
in the variously metaphorphosed adult. Thus laid down
the nerve-cell keeps in touch with the remainder of that
segment to which it belongs in the body by virtue of the
enormous power of individual growth of its cell-branches.
It does not, as in the young muscle-cell, keep pace with the
extension of the body segment in any one direction by means
of a repeated fission of nucleus and reproduction, and the
forming of a chain of muscle-cells, more or less nutritively
independent. ‘To trace such a line of units, although they
are closely connected by blood-relationship, is impossible
with certainty by any known method. But the nerve-
cells remains securely anchored in their original segmental
positions, both inside the spinal cord and outside it, in
the row of segmentally-arranged spinal ganglia. Growing
out from the nerve-cells their nerve-fibres wander dis-
tally into skin and muscles, and the main difficulty in
tracing the actual limits of the metameres in these last,
is that the elements of the adjacent metameres certainly
considerably intermingle. So it 1s also with the ramifica-

RELATION BETWEEN STRUCTURE AND FUNCTION. 7

tion of the nerve-cell branches within the brain and spinal
cord themselves.

It may be conceded, therefore, that the nerve-cell and
fibre are trustworthy guides to the plan of the segmental
architecture of the body and of the hmb. Various other
indices have been used, such as the bones, muscles, &c.
The former of these led to the view (Goodsir) that the
number of digits of the limb told the number of meta-
meres in the limb. As to the muscles, the examination
by the nerve-distribution method shows that even the
most homogeneous-seeming muscle may be traceable to
belong to two and three segments, and be combined from
elements of each. It is apoint upon which the embryo-
logist is undecided whether the myotoms grow out into
the limb-bud; so that embryology is evidently quite
unable to say even should the myotoms be proven, as is
likely, to gYOW out into the limb, and so give rise to its
musculature, what particular part of the limb musculature
is the outcome of any one particular myotom.

What is the result of the analysis by the nerve-distribu-
tion method? As regards the muscular part of the
metameres some of the results are indicated in the
accompanying synopses. The synopses include the
range of individual variation, so that although the e.g.
flexor sublimis digitorum muscle is considered as quadri-
segomental, 7.e., belonging to four body segments of the
brachial region in the species, it in any individual of the
species will belong only to three. To this matter of
individual variation I will return later.

The topographical representation of the various elemental
movements of the limb in the chain of segments of the
spinal cord is always arranged in a definitely ordered
sequence. ‘This sequence is exhibited in the accompany-
ing diagram :—

8 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

x | 1x [vill] vil} vi
Protraction of shoulder- - -

| Abduction of shoulder - - -

Vv
x
x
Outward rotation of shoulder - x
x

Flexion of elbow - - - + -
Supination of forearm - - - ?
Adduction of shoulder - - - x x

Radial abduction of wrist - -

x

Extension of wrist - - - -

Inward rotation of shoulder -

Re sONe Ok ORR. aR KS ee.

Extension of fingers- - - -

Extension of elbow - - - -

KS)
"~ .

Flexion of wrist - - - - -

Xx

Retraction of shoulder - - -| X

RS)

Pronation of forearm

Te Soe ORS OG ei Ce IK OK OG
>
>

Flexion of fingers- - - - -| ?

xK ©
Ms NM SX “XK Be. oe, OS OX

Interosseous flexion of fingers -

Adduction of fingers - Fibs agkhee

MS IK ARK TS Oe RE ie a OG BE, EK

Abduction of fingers- - - -| xX

Certain movements of the hmb are not seen at all in
the stimulation of individual motor roots: e.g., in many
monkeys no root gives extension of the pollex, and in
some monkeys no root gives extension of the wrist, and I
have met with an individual in which no root evoked
extension of the fingers.

RELATION BETWEEN STRUCTURE AND FUNCTION. 9

The range of the spinal innervation of the skin and the
muscles covering, and moved by and moving, the joints of
the upper limb, is shown in the accompanying conspectus.
The sensory innervation of the deep structures may be
looked on as similar in segmental range to the motor
innervation of the muscles.

if horax.

ive ti 1, 1
Number of spinal |_____».___
merve-root - -| xii| x1{ x | 1x |vll1| Vil} vi

an hae!
skin % Mellie, 5 * * * * * *
Shout
muscle - NOG Ga OG. | Oe I
skin- 2 * * * * * *
muscle - xX|xX |X| xX
Elbow
flexors - CAL CAS a

extensors ><a > <a 2a ae

skin - -
muscle - x SOX
Wrist
6 Silke

x
x

x)

x

x
~

flexors -

extensors DIES De SM

skin - -

x
x
~

muscle -

Fingers

x
x

flexors -

~~

x

x
~D

extensors

10 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

I have given recently a synopsis of the spinal nerve-
supply of the muscles of the upper limb, which shows
that the muscular tissue of the limb is arranged in a
number of rays, a ray for each metamer contributing to
the limb. It shows that of these rays the ones which
lie tailmost in the fore-and-aft series are the longest, that
is, extend to the extreme free apex of the limb, whereas
the foremost, the most rostral, pass only as far as the
upper arm, the next only as far as the elbow, the next
only as far as the wrist. The four last, or most aboral rays,
all contribute to the musculature of the hand in Macacus
rhems, the common Rhesus monkey.

But when we come to enquire how these units of the
segmental architecture of the limbs, these muscular rays,
are related to the physiological or functional units of the
limb-musculature, it is at once obvious that the extent
and boundaries of the two do not coincide. The individual
muscles of the limb are functional elements of its structure
as a physiological machine. But each of these functional
elements is compounded of portions of several rays or
myotoms. Moreover, the boundaries between the
myotoms do not correspond with the intervals of muscles,
or even between muscle-groups.

The same want of segmental separation 1s evident when
we examine the location of the nerve-cells which innervate
the individual muscles, Hach of the segmental spinal
nerves which supplies the musculature of the limb leaves
the spinal cord in the form of a row of rootlets, which
together build up the motor-root of the nerve. Excitation
by electric currents of anyone of these rootlets or fila-
ments, that contribute to the formation of the motor-root,
produces a movement in the limb which resembles closely
the movement produced by excitation of the whole root.
The same muscles are thrown into contraction as when

RELATION BETWEEN STRUCTURE AND FUNCTION. 11

the whole root is excited; but the action of each muscle
is feebler. The collection of nerve-fibres contained in the
rootlet is therefore, in miniature, a collection like that
contained in the whole root. This fact must be taken
together with the fact that, as can be shown by degeneration
experiments, the fibres contained in each rootlet repre-
sent the nerve-fibre processes, or axis cylinder processes,
of the motor nerve-cells lying in that particular transverse
plane or level of the spinal cord, from which the rootlet
makes its emergence. ‘The two facts together show that
the nerve-cells which innervate the different muscles
belonging to any one myotom in the limb, lie commingled
together, so that any one section through the spinal
cord, taken at right angles to its length, will meet nerve-
cells belonging to all the muscles represented in that

segment. This is the same as saying that the nerve-cells

belonging to any one limb-muscle, lie not gathered
together at any one particular level of the spinal segment,
but scattered throughout its length. And, inasmuch as
each muscle is innervated from several segments, the
nerve-cells for each muscle are scattered in a continuous
series through the length of a series of spinal segments;
and throughout this extent are commingled with the cells
of a great number—in some cases as many as forty and
more—of other muscles.

It is, therefore, evident how it comes about that no
traumatic injury of the spinal cord can ever paralyse a
single muscle alone and apart from its fellows. Even
the severance of a whole single motor nerve-root cannot
paralyse a single limb-muscle; the effect of such an injury
is to partially impair a large number of the muscles.

It has been urged by many observers that the motor
nerve-fibres gathered together in each motor spinal-root
are a collection connected for functional co-operation.

12. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

These writers attach, therefore, a clear and simple func-
tional meaning to this collection which composes the
spinal root. They point to the fact that stimulation of
this root or that root produces a movement of the limb
which is of some definite kind, such as a flexion of the
elbow, flexion of the fingers, &. The movement evoked
they consider one which indicates that the nerve-fibres
associated together in the motor root are associated there
in order to, by acting contemporaneously, execute a highly
co-ordinate functional synergy of muscles. By synergetic
muscles are understood those muscles which, in natural
movements, contract harmoniously together to execute
a movement. I find nothing which is really valid evi-
dence for this view and much to the contrary. The
mere inspection of a movement of the limb without
further analysis of 1t 1s very Insecure a guide by which to

judge of its natural character. The joints and muscles of |

the limb have been evolved contemporaneously and
together in the course of the history of the individual and
of the species. No muscle can, therefore, be thrown into
-action which will move the limb in any which is an
unnatural direction. Our knowledge of the synthesis of
the natural movements of the body out of the contractions
of the individual muscles is extremely meagre. The
names given, for instance, to many of the muscles of the
human arm are misleading rather than helpful as regards
the movements they produce. The resolution of the com-
bined movements into their components require data
hardly obtainable, because requiring isolated contraction
of individual muscles in life, and the alteration of the
lengths and tensions and elasticities of the muscles after
death, precludes the possibility of dealing with them even
approximately accurately by examination in the dissecting-
room. Thus few realise that the gastrocnemius muscle

RELATION BETWEEN STRUCTURE AND FUNCTION. 18

is a flexor of the hip, although this can, as a fact, be experi-
mentally shown; and the mathematical treatment of
these problems is far from easy.

Against the view that any one spinal motor - root
represents the motor spinal nerve cells concerned in some
particular highly co-ordinate functional synergy, rise a
number of facts. These I have detailed elsewhere.
Among them are the sudden change which in some
motor-roots occurs as the stimulation of the indi-
vidual rootlets composing the root is undertaken in
serial order; the upper filaments of the root producing one
movement, the lower series another movement; the upper
filaments of the root produce a movement like that
_ produced by the filaments of the root next above, the
lower like that of the root next below. Here, then, the
collection of the root would contain two functional groups
representing two synergies, and the upper one would be
simply a replica of that of the functional synergy of the
root next above, and the lower that of the root next
below. Then the combination of a piece of the diaphragm
muscle with supination of the wrist, of flexion of the wrist
with extension of the fingers, are bizarre effects unknown
in any reflex movement, or movement produced by
excitation of the brain. The combination of supination
at the wrist with flexion of the hand, is of itself an unusual
movement, that usually requires some attention for its
execution.

The degeneration experiments show that in some
muscles the number of motor nerve-fibres given by a
spinal-root to a muscle is too small to evoke from the
muscle any contraction at all obvious to inspection.
Cases occur where a limb muscle receive one, two, three,
four, or five motor fibres from a particular root; allow to
each of these motor nerve-fibres a dozen muscle-fibves, it

14 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

is easy to understand that sixty muscle-fibres scattered in
a muscle consisting of many thousands may cause no per-
ceptible tightening of the tendon; they may simply stretch
or compress adjoining inactive and elastic fibres. The
degeneration of these few fibres I look upon as strong
evidence of the morphological character of the overlap ;
the paucity of the fibres is one of the many facts which
indicate that the distribution of the motor-roots is arranged
on a segmental plan in accordance with the terms of a
bequest dating back to a time when the present environ-
ment of the limb, especially in its Mammalian form, had
no preponderant weight in the shaping thereof. As re-
gards functional value this character of the Mammalian
limb is on a par with details of structure which are not
specific, and, therefore, with other details. of structure
outside these immediately acquired by the species, not to
be considered as of necessity of functional importance.
The functional use of the contribution of one or two nerve-
fibres to a muscle requiring hundreds is difficult to see;
the probability of the occurrence of such poverty-stricken
contributions is, on the view of the morphological neces-
sity of the ray-arrangement of the limb musculature, so
high as to be only what might have been expected from
theoretical considerations. The fact that certain of the
motor-roots of the limb contribute fibres to the innerva-
tion of certain muscles in such scanty number as to be
ineffective for movement is a further argument for the
morphological rather than functional character of the
motor-root distribution in the Mammalian linb. A number
of motor-fibres, too small to evoke appreciable movement
-in a muscle when excited electrically, will hardly be effective
for movement under the action of the will.

Strongly militant against the functional theory of the
motor-root is the frequent occurrence and wide range

RELATION BETWEEN STRUCTURE AND FUNCTION. 15

of individual variation in the movements produced by
excitation of the roots, and in the distribution of the
nerve-fibres composing a root to the muscles of the limb.

In experiments on the plexuses, it is rare to find two
consecutive individuals of the same species that possess
a similar root distribution. The plexus in any species
requires for description to be grouped into classes. For
my purpose I have found two classes enough, a class of
individuals in which the plexus is pre-fixed and a class in
which it is post-fixed. Thus, in some individuals, the
supinator brevis is innervated from the 6th and 5th
cervical nerves, in others from the 6th and 7th. In the
former case the plexus is of pre-fixed type, and when one
muscle or one part of the plexus is pre-fixed, all the rest
of it is pre-fixed, and conversely, when one part is post-
fixed, all is post-fixed. In the second instance given
above, the supinator brevis is post-fixed, that is, its nerve-
cells are fixed in segments further aboral in the series
than in the individuals with pre-fixed plexuses. Presum-
ably the muscle itself therefore is built of myotoms more
aboral in the series than in the muscle in the pre-fixed
individuals. But instead of two classes, there might be
made a very large number of classes were it worth while
to separately distinguish them, for between the extreme
cases of post-fixed type and the extreme examples of
pre-fixed type extend a numerous series of intermediate
individuals ; in fact, any and every intermediate degree of
type seems to exist. The extreme of post-fixed type
passes over into the comparatively rare individual with an
additional segment altogether, e.g., the cat with fourteen
ribs, which is not at allan uncommon find in the laboratory.
Never a year passes without our meeting with four or five
such. This frequency of individual variations exhibits in an
almost ludicrous aspect the view that each motor spinal

16 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

root is a collection gathered together for a functional
purpose, and representing a particular highly co-ordinated
movement of the limb. Contrary to any such hypothesis,
it shews that for the functional mechanism of the limb
it is immaterial whether this or that particular group
of muscles is represented in a particular spinal root or
not represented in it. The absolute segmental level is
variable over the range of nearly a whole segment’s
length; the relatwe segmental position is, however,
inviolably preserved. Thus flexor carpi uluaris may get
fibres from as high as the 6th cervical segment, or it may
not, but it never extends so well forward, so far rostrally,
as flexor carpi radialis, and this latter never extends so
far aborally as flexor carpi uluaris, even when it is most
post-fixed; that is the region of out-flow for the spinal
nerve-fibres to flexor carpi uluaris lies rather more aborally
than that of the out-flow of flexor carpi radialis, and this
mutual segmental position of the muscles is maintained
whether the plexus be post-fixed or pre-fixed.

The idea at root of the supposition, that the motor-root
is a functional collection of nerve-fibres, seems to be that
juxta-position in space is advantageous for co-ordination
of action. The material of support for this idea is very
slender, and there are many facts which show that mere
spatial separation offers no difficulty for the physiological
mechanism of co-ordinate and synchronous activity. The
great respiratory centre in the spinal bulb, and the various
subsidiary spinal respiratory centres, lie very far remote
one from another. Yet how accurately do they harmonise
in their activity? Their co-ordination consists rather in
the fact that they are all chemically tuned, so to say, to
the same pitch or note of stimulus. They differ from all
the other bulbo-spinal centres, and agree among themselves
in their peculiar sensitivity to the stimulus of venous

RELATION BETWEEN STRUCTURE AND FUNCTION. 17

blood. So also in the case of the individual spinal nerve-
cells, innervating a particular muscle, we suppose, that
their co-ordinate action and synchronous discharge depends.
rather on some physiological quality common to all which
cannot be accorded to them by mere spatial arrangement.

A question that obviously presses for answer, 1s whether
when a muscle contracts either reflexly or voluntarily, all
the parts of it innervated from all the spinal roots con-
tract together, or whether some parts of it contract for
some movements, and others for other movements, and
so on, and so whether the segmentally different parts of
the muscle are used severally for different functional
actions. In reflex movements, and in movements evoked
from the cortex of the brain, we can say that the discharge
of motor impulses to a pluri-segmental muscle of the fore-
arm, or wrist, or hand, is always pluri-segmental, and
involves the whole length of the serial group of intra-spinal
nerve-cells which innervates the muscle extending through
its full number of segments of the cord. The afferent and
cerebral channels of the cord treat the pluri-segmental
motor stations or nuclei of these limb muscles as entities
of homogeneous structure, as in fact, physiological units.

This serves to emphasise the physiological homogeneity
of limb-muscle and nerve-trunk, and the physiological
heterogeneity, in spite of morphological unity, of the
spinal nerves-root in the limb-region of the body.
The spinal nerve-roots of the thoracic region are,
from the physiological point of view, less heterogeneous.
The peripheral nerve-trunk is the physiological collection
of nerve-fibres, e.g., flexors collected together, vaso-
dilators, included with motors to muscles, &c. The nerve-
root is the morphological collection, it contains com-
mingled into one such heterogeneities as adductors of the
hallux with protrusor pelvic muscles.

18 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Similarly with the skin, the median nerve-trunk
supplies a patch of the palm that has obviously functional
unity, but Ist thoracic nerve-root supplies such incon-
gruities as the front of the little finger and of half the
ring finger, together with the tip of the point of the
elbow.

It is the formation of functional collections of nerve-
fibres (peripheral nerve-trunks) out of morphological
collections (nerve-roots) which is the meaning of the

interlacements of adjacent spinal segmental nerves in the

limb plexuses. The reply to the frequently askea question
as to the explanation of the distribution of the spinal
nerves of the brachial and pelvic limbs by plexuses, while
the spinal nerves of the trunk region are not distributed
by plexuses, is in my opinion as follows. In the trunk
region the innervation of the muscles of the skin 1s, as
regards the distribution in them of the segmental nerves,
a system of comparatively slight overlap: the peripheral
territory of each segmental nerve—especially each motor
territory—is confluent with, but does not mingle nearly
so widely with the neighbour territories as in the limb

regions. That is, in other words, each several area

of skin and of muscle, especially of the latter, has in
either of the limbs a more pluri-segmental spinal innerva-
tion than a comparable area in the trunk. The anatomical
mode of innervating a definite area of tissue is, as we know,

by means of collecting tle nerve-fibres for the region into ©

a nerve-trunk; where the innervation is pluri-segmental
the nerve-trunk will naturally be combined from
components of several segmental nerves—where several
such areas co-exist several pluri-segmental nerve-trunks
will be formed and the separate segmental nerves will be
split up into components, which become redistributed in
the combinations which constitute the pluri-segmental

RELATION BETWEEN STRUCTURE AND FUNCTION. 19

nerve-trunks of the region—as, for instance, in the brachial
reigon. The brachial and lumbo-sacral plexuses are an
anatomical result of the greater degree of overlap, especially
in the distribution of the motor part of their spinal nerves,
obtaining in the limbs as compared with other, e.g., the
trunk regions of the body.

Enough has been said, I imagine, to let us see how little
light is thrown upon the function and uses of the limbs by
study of their segmental architecture. It is especially when
the anatomist confines his attention to minute examination
of one limited group or type, that he seems to become
most prone to dogmatise concerning function, and his
arguments have to be received, unless checked by actual
experiment, with the extremest caution. In a certain
sense, it would seem that the creative power of nature can
mould with such superfluity of resource the, to us, who
cannot construct it, pricelessly valuable material that
heredity and life bequeath, that she has various ways
open for compassing the same required end. ‘Thus a seg-
ment more or a segment less offers little difficulty to her.
The individual with a segment less or segment more
betrays neither his defect nor his surplus by any trace of
disturbance of function; his idiosyncrasy goes unsuspected
either by himself or his neighbours, physiologists and
morphologists though they may be. When the “‘how”’ of
the processes of function and of structure is nearer answer
than at the present time, this may appear to us less strange
than now. But for the present it does seem clear that the
aws of morphological structure and of physiological struc-
ture, though they must be fundamentally correlated, are
much more different than many are accustomed to
think of them and say. The width of the chasm between
the two is probably best felt by the zoologist who holds in
view in his mind’s eye the vast range of comparative

20 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

structure in animal forms, and by the physiologist who,
studying the body in its aspect as a mechanism
for liberating the locked energy of chemical mole-
cules in forms of material, and perhaps of mental,
- life finds an extravagance of material and a riot of
design that baffles his description almost as much
as his comprehension. But he always remembers
that his question is not why, but how. When
Helmholtz, after his investigation of the optics of
the human eye-ball, drew attention to the mani-
fold optical defects in that structure, and wrote that
an optician would discharge a workman for producing
so optically imperfect an instrument, it appeared to some
that such a criticism was impious in character. The
attribution of impiety to such expression shews how hable
to confusion are the two lines of enquiry of the why and
how. As Helmholtz himself points out in the most
beautiful of passages, so inconceivably admirable is the
wonderful mechanism of the nerves and brain upon which
the optic parts of the eye-ball throw their relatively
imperfect pictures of the visible world, that despite these
defects, the sense of vision results, with all its marvels,
and builds the chiefest basis and scaffold for the uprearing
of the indescribable edifice of the human mind.

21

TWELFTH ANNUAL REPORT of the LIVERPOOL
MARINE BIOLOGY COMMITTED and their
BIOLOGICAL STATION at PORT ERIN.

By Professor W. sae HERDMAN, IDSGa5 LE leictsis

THE Report affords an opportunity of laying annually
before the Biological Society, our subscribers, and the
larger public of Liverpool and the neighbourhood, not only
a statement of the statistics of the Biolovical Station
during the year, and a record of the work done, but also
an outline of such plans for the future, extensions of the
work, and promising investigations as seem to the Director
worthy of being published, in the interests of Biological
science, in order that some possibly who are able may
follow up the suggestions, and that others who can afford
it may, if they care, provide funds to enable*%special pieces
of work to be undertaken.

The year’s work will, as usual, be reported upon first,
and then certain proposed investigations and more general
matters will be discussed (see p. 43).

Tur STATION RECORD. |

During the past year the following naturalists have
worked at the Station, in addition to the Curator, who
has been constantly in attendance :—

DATE. NAME. WORK.
January Mr. 1. C. Thompson, Liverpool ... ..- Copepoda.

-—- Professor W. A. Herdman, Liverpool ... General.

os Dr. Christophers, Univ. College”*... pe cremeral.

* Univ. College after a name indicates that that worker made use of the
“table” secured by the College Council,

22 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

DATE.
March.

August.

September.

October.
December.

NAME.
Mr. I. C. Thompson
Prof. W. A. Herdman
Mr. F. J. Cole, Univ. College
Mr. R. L. Ascroft, Lytham
Mr. F. J. Cole, Univ. College
Mr. A. R. Jackson, Univ. College
Mr. J. C. Mann, Univ. College
Prof. W. A. Herdman
Mr. Mundy, Owens College
Mr. I. C. Thompson
Mr. A. O. Walker, Colwyn oar
Rey. L. J. Shackleford, Cline
Rev. T. S. Lea, Liverpool ...
Mr. F. J. Cole, Univ. College
Rev. T. S. Lea
Prof. W. A. Herdman
Mr. I. C. Thompson
Mir *A. O; Walker =.
Mr. J. A. Clubb, Liverpool Mido

‘Mr. R.. L. Ascroft ... a.
Mr. C. Crossland, Clare College, ea shag

Mr. R. L. Ascroft ...

Rev. L. J. Shackleford

Mr. H. Yates, Manchester...
Rev. L. J. Shackleford

Mr. I. C. Thompson

Prof. W. A. Herdman

Mr. F. J. Cole, Univ. College
Mr. I. C. Thompson

Prof. W. A. Herdman

WORK.
Copepoda,

Comp. Ascidians.
Comp. Ascidians.
Fishes.

Comp. Ascidians
Arachnida.
General.

Comp. Ascidians.
General.
Copepoda.
Amphipoda.
Mollusca.

Alge & photography
Comp. Ascidians.
Alge, &e.

Comp. Ascidians.
Copepoda.
Amphipoda.
Actinia.

Fishes.

General.

Fishes.

Mollusca.

Poe & Plankton.

Mollusca.
Copepoda.
General.

Comp. Ascidians.
General.

General.

The work of a number of these Naturalists will be

found discussed further on in this report.

THE CURATOR’S REPORT.

Mr. H. C. Chadwick entered upon his duties early in

January, 1898.

Since then he has kept the Station in

excellent condition, has added greatly to the attractiveness

of the Aquarium, has carefully attended to the wants of .

a a

a

MARINE BIOLOGICAL STATION AT PORT ERIN. 23

workers in the laboratory, and has done a considerable
amount of collecting on the shores and in the bay. A most
effective addition to the collecting apparatus which can be
worked from a rowing boat in the bay is a small iron
trawl of 44 feet beam, and fitted with the cod-end of a
shrimp net. This is lighter to pull than a dredge, sweeps
a greater area, and gives better results.

The Lords Commissioners of Her Majesty’s Treasury
have kindly presented to the library of the Station the ©
three large quarto volumes containing the well-known
elaborate Report upon the “ Challenger’? Amphipoda by
the Rey. T. R. R. Stebbing, F.R.S., a classic work in
marine biology.

Mr. Chadwick reports as follows :—

“The Port Erin Biological Station has been open every
week day from January 24th to the present. The Labora-
tory, together with the instruments, dredges, and other
implements has been kept in a clean and efficient condition.
A few additions have been made to the stock of reagents,
and the supply of all has been kept up to the requirements
of workers. A few additions, in the shape of reprints
and one volume (purchased) have been made to the library.
To all donors an acknowledgement was promptly sent.

‘‘ Meteorological observations have been taken and care-
fully recorded twice almost every day, the very occasional ©
omissions being caused by the Curator’s absence on a
dredging excursion.

“In January, the temperature of the sea was almost
uniformly 2° higher than that of the air, while in February
it averaged 4° to 5° higher. In March, it averaged about
3° higher, and in April, 1° higher. From the beginning
of May to the end of September, the temperatures of air
and sea averaged the same. ‘The greatest disparities
occurredin May, when, during the afternoon of the 13th,

e

24 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the temperature of the sea was 11° higher than that of the
air, and during the afternoon of September 5th, when the
temperature of the sea was 12° less than that of the air.
The density of the sea-water has shown very little
variation, and only twice have I noticed a slight loss of
density to occur immediately after a few hours rain. The
average may be stated as 1:026.

Fig. 1. The ‘‘ Shellbend” punt on the shore—starting for a
“plankton” trip. [From a photo by Mr. Edwin Thomyson. ]

“The ‘ Shellbend’ folding boat has been frequently used,
and is still in a thoroughly seaworthy condition, though
beginning to show signs of wear. In launching and
housing the boat residents and many visitors have
rendered willing assistance.

‘Tn spite of much rough weather, over fifty tow-nettings
have been taken and carefully preserved. The Curator’s

MARINE BIOLOGICAL STATION AT PORT ERIN. 25

time has been so fully occupied with other duties that he
has not yet been able to carry out the projected careful
examination of these week by week. Sagitta appeared in
ereat abundance on June 2ist, but has occurred only
sparingly since. Plewrobrachia has occurred in all the
gatherings from early spring onwards, and the same remark
applies to Appendicularia.

“The Scyphomedusze have been very sparingly repre-
sented this year. Several trips were taken in the ‘Shellbend’
in quest of Aurelia aurita, but none were obtained. A
large number were, however, cast ashore on one day early
in July. About twenty large specimens of Rhizostoma
pulmo were cast ashore early in March, in the gastric
canals of each of which were a nwuber of specimens of
Hyperia galba, and one living speciien was seen in the
bay on October 12th. Cyanea capillata and Chrysaora
isosceles have not. come under the Curator’s notice.

‘““Hleven dredging excursions have been undertaken in a
small boat, and hauls taken in various parts of the bay,
outside as well as inside the break-water. In one such
excursion the following species were taken :—Clytia
johnston, Obelaa geniculata, Asterias rubens, Astropecten
wrregularis, Ophiura ciliaris, O. albida, Echinus esculentus,
EH. miliaris, Nereis sp., Portunus pusillus, P. puber,
Stenorhyncus rostratus, Hupagurus bernhardus, E.
prideauxw with Adamsia palliata, Crangon vulgaris,
Virbius varians, Galuina farram, Facelina drwmmondi,
Aplysia punctata, Rissoa parva, var. mterrupta, Phasia-
nella pullus, Seprola sp., egg capsules of Loligo forbesz,
with developing embryos.

“The frequent occurrence of rough sett has prevented
the Curator from gaining more than a general idea of the
nature of the bottom of the bay beyond the lowest tide
mark. ‘The shoreward border of the great .sea-weed bed,

26 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

which trends in a south-westerly direction from the spot
where the drainage of Port Erin is discharged, has,
however, been traced.

“Shore collecting has been carried on at intervals all
through the year, and much knowledge gained of the
precise localities in which many species may be found.

‘On August 15th the buoy at the end of the breakwater
was brought ashore for its annual cleaning, and with the
assistance of Mr. C. Crossland of Clare College, Cambridge,
the Curator took from its flat bottom the following
species :—

‘““Sycon compressum, Bougainvillia ramosa, Coryne sp.,
Tubularia larynx, Hudendrium ramosum, Obelia genicu-
lata, Clytia johnstom, Sertularella rugosa, Nereis sp.,
Terebella sp., Sabella sp., a Nemertine, Bicellaria ciliata,
Scrupocellaria sp., Membranipora, several Amphipods and
Isopods, Nymphon gracile, Doto fragilis, Ciona mtesti-
nalis (very abundant), Ascidia mentula, Styela grossularia
(abundant), Corella parallelogramma, and at least half-a-
dozen as yet undetermined species. Such a large
assemblage of animals, over thirty species, formed a
striking commentary on the efficacy of the anti-fouling
paint with which the buoy had been covered, and which,
the harbour master said, had been guaranteed by the
makers to poison everything.

On November 2nd there was another incursion of the
Amphipod Orchestia gammarellus from the shore. As
on previous occasions they swarmed all over the laboratory.

‘The number of visitors to the Aquarium again shows a
substantial increase, over 500 having paid for admission.
The constant care which its successful maintenance has
entailed has been repaid by the interest shown by the
visitors, and by the favourable comments of many who
had visited it in former years. A considerable proportion

MARINE BIOLOGICAL STATION AT PORT ERIN. 27

of the visitors were families of children who were brought
by their parents to learn something about the structure
and habits of the animals seen in their rambles amongst
the rocks on the shore. The fish-hatching tanks have
attracted a great deal of attention, and the many questions
asked by the visitors—questions based on information
gained from newspapers and magazines—showed that the
interest in fish-hatching is becoming wide-spread.

“We also had in July a visit from a number of the
pupils of the Arnott Street Board School in Liverpool,
the party being under the guidance of Rev. 'T. 8. Lea.
The Curator hopes before next season to arrange a series
of tanks, for workers at the Station, through which sea-
water will constantly circulate.

“The publication of a price list of specimens has not as
yet had very much result, comparatively few orders having
been received. The list, thoroughly revised and with
prices lowered as much as possible, is now appended to
this report.”

ADDITION TO THE LABORATORY.

During the past year we have added very considerably
to the accommodation for students at the Biological
Station by placing a wooden staircase and a new upper
floor in the laboratory. This upper chamber is sub-divided
by partial wooden partitions into five alcoves, each
having a small window in the roof, and each containing
a fixed work table for one student, measuring over 8 feet
by 3 feet. This increase of accommodation permits the
Committee now to carry out a plan they have long had in
view, viz., to arrange an

HASTER STUDENTS’ Parry.
It is proposed that any persons who desire to join as
students should pay (in addition to their own expenses)

—————iL——<o— “~"—
[roy “g *y aay Aq ogoyd vw wo1g] ‘vas oy} Woy ‘sepry MOL 4e ‘SHUIPUNOLINS s}I PUR UOTZRYg [vOIBO[oTg ULI Wog *z ‘Sty

SOCIETY.

.

TRANSACTIONS LIVERPOOL BIOLOGICAL

28

MARINE BIOLOGICAL STATION AT PORT ERIN, 29

a small fee to the funds of the institution. * Dredg-
ing, tow-netting, and shore collecting expeditions
will be organised on certain of the days, accord-
ing to tides and weather; and at other times there
will be occasional lectures and demonstrations in the
Biological Station, and opportunities for laboratory work.
Several members of the Committee have promised to join
the party, and the scheme offers a very unusual oppor-
tunity for collecting and studying in the company of
specialists which will probably be of great service to
junior students of marine biology and to some amateurs.

DREDGING EXPEDITIONS.

In addition to frequent dredging and trawling from
small boats in the bay, the following expeditions in steam
trawlers have been organised :—

I. April 9th, 1898. In Fisheries steamer “ John
Fell’’—
1. Two miles off Fleshwick, 20 faths. (2 hauls
of shank net).
2. Two miles off Dalby, 35 faths., bottom
mud (with shank net). -
3. ''wo-and-a-half miles off Dalby, 40 faths.,
bottom mud (with fish trawl).
ie Apr ith, 1898. In Fisheries steamer ‘‘ John
Fell ’>—
1. One to two miles off Dalby, 20-35 fathoms.
2. Between Bradda Head and Calf Sound, 17
faths.
Ill. April 20th, 1898. In steam trawler ‘ Tudor
Prince ’’—
From three to five miles off Entrance to
Glen Meay, 22 to 30 faths.

* A cireular, giving all particulars, will be issued. Apply to the Curator.

30 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

IV. July 16th, 1898. In steam trawler ‘‘ Rose Ann,”’
when we dredged and trawled at the following
localities :—

2s
EIGER POOL

ay
HSN ‘it \
{ EH HN i}

Fig. 3. Plan of the L.M.B.C. District.

1. T'wo miles south of Spanish Head, 17 faths.,
a neritis bottom.

2. Four miles east of Port St. Mary, 18 faths.,
many Ophiuroids.

3. Four miles east of St. Ann’s Head, 20
faths., gravel bottom.

4. Three-quarters of a mile east of St. Ann’s
Head, 13 faths., melobesia bottom, drag-
ging inwards to Alge and Polyzoa, with
enormous masses of Perophora (chiefly on
Vesicularva spinosa) and other Ascidians. In
this last haul, 14 species of Tunicata were

_ MARINE BIOLOGICAL STATION AT PORT ERIN. 81

obtained, as follows:—Perophora lstert,
Clavelina lepadiformis, Ascidia mentula,
A. scabra, Corella parallelogramma, Mol-
gula sp., Molg. sp., Polycarpa pomaria,
P.comata, Styelopsis grossularia, Amarou-
coum proliferum, Aplidiwm sp., Diplosoma
sp., and Botryllovdes sp.

We still have living healthily in the tanks specimens of
the following moderately deep-water forms obtained at
one of the dredging expeditions last summer :—Stichaster
roseus, Porana pulvillus, Venus verrucosa, Pectunculus
glycumerts, and Aporrhais pes-pelicant.

The following rare or interesting animals have been
kept in the tanks of the Aquarium house during the
year :—

Zoothamnion dichotomum.

Hphyre of Aurelia aurita.

Lineus gesserensis.

Carinella annulata.

Phyllodoce laminata.

Cucumaria hyndmanni.

Hyperva galba.

Eggs of Loligo forbesv.

The sea slugs Galvina cingulata and Holis farram.

Lepadogaster decandollit.

The Amphipod T’riteta gibbosa, infesting colon-
ies of Amaroucium argus.

A specimen of Hupagurus prideauxw (the hermit crab),
which had Adamsia palliata (the cloak sea-anemone) on its
shell, after being kept for some time, adopted a new shell—
the Adamsia accompanying it.

Early in July great shoals of young Gadus virens (the
saithe) and Pleuronectes platessa (the plaice) were seen
close to the shore, in front of the laboratory. They were

32 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

feeding upon the “fairy shrimp,” Mysis inermis, which
swarmed at the time in countless thousands. Mr. Andrew
Scott finds, on the Lancashire coast, that young Herrings,
2 to 8 inches long, feed largely upon Mysis. There can
be no doubt that this Crustacean 1s an important fish food.

Notes oN WorRK DONE AT THE STATION.

Mr. J. A. CLUBB, M.Sc. (Vict.), is now carrying out his
investigations on the structure and variations of sea
anemones upon material from Port Erin. He spent
some time at the station in summer collecting material
for his work, the first part of which has just been published
in our twelfth volume.

A quotation from Mr. Clubb’s report to me runs :—

‘“‘T had a very good collecting time during the remainder
of my stay at Port Erin. We dredged Adamsia palliata,
obtaining about half-a-dozen good specimens, and I
succeeded in fixing three of them well expanded. I did
not, however, get Corynactis viridis, although out dredging
for it two or three times. I got a number of the littoral
forms (five different species), and have fixed them all for
dissection and histological work.”

Mr. J. NEwton Coomsgs, late Chairman of the Sheffield
School Board, has been supplied frequently with gather-
ings of diatoms from the surface during the spring and
early summer. From this material he has been making
observations upon the life history and reproduction. and
his results, he informs me, are nearly ready for
publication.

Mr. A. O. WALKER sends me the following ‘‘ Report on
Malacostraca in 1898”’ as the result of his work this year :—

“There has been a remarkable scarcity of Amphipoda
this summer on the coasts of Liverpool Bay within a mile
of the shore. At Bull Bay, Anglesey, in June; Port Erin

MARINE BIOLOGICAL STATION AT PORT ERIN. 33

in July; and Colwyn Bay in September, very few
individuals were taken in the bottom tow-net. It is
possible that this may partly account for the small takes
of fish at Rhos Weir this summer.

“The following are the more noteworthy Crustacea
taken during the past season. :—

‘“‘ Mysis longicornis, M. Edw. [G. O. Sars, Middelhavets
Mysider, p. 22 (separate copy) pls. 1x and x.]

“This species, hitherto recorded only from the Bay of
Naples, may be at once distinguished from all the other
British species of the old genus Mysis,except M. (Neonvysts)
vulgaris, J. V. Thompson, by the absence of a notch, or
cleft, at the end of the telson, and from that species by
its telson haying the end rounded, and the margins
furnished with closely set setze of unequal length. On the
other hand, the resemblance of the telson and antennal
scales to the genus Leptomysis is remarkable. The female
of this species is practically indistinguishable from the
same sex in Leptomysis aptops, G. O. Sars, except by the
length of the spine on each side of the two small central
spines at the apex of the telson. These in M. longicornis
are but little longer than the other long spines on the
margin, while in L. apiops they are quite twice as long.
The resemblance extends to the curious trigonate points
of the spines round the end of the telson, which I have
not observed in any Crustacean except these two. A new
genus is required for this species, the characters of which
do not agree with any of the existing genera of Myside.

“One female was taken in the bottom tow-net at Bull
Bay, June 10th, 1898, in 20 fath. Length 6 mm.

“Cumella pygmea, G. O. Sars. Several females and
three males taken at the same time and place as the last.
Not previously recorded in Liverpool Bay.

“Argissa hamatypes (Norman); = Syrrhoé hamatipes,

34 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Norman, Brit. Assoc. Report on Shetland Dredgings,
1868, p. 279; = Argissa typica, Boeck, Crust. Amph.
Bor. et Arct. 1870.

‘‘A single male specimen was taken in the bottom tow-
net at Colwyn Bay in 23 fath., Sept. 13th, 1898.
I have Dr. A. M. Norman’s authority for saying that
his species is identical with Boeck’s, and I have seen
the type specimen. Previously recorded from the Isle of
Man coast in 12 fath.

“Gammarus duebem, Lilljeborg.

‘“‘ A single specimen, dredged off Glen Meay, about four
miles from land in 22—380 fath., April 20th, 1898.

“There is, perhaps, no aquatic animal which appears to
be able to exist under such different conditions as this
species. I have specimens collected by Dr. R. F. Scharff,
in Lough Doon, Co. Kerry, ‘‘ about 1,000 ft. above sea-
level;’? Mr. Thos. Scott records 1t from Loch Ruan, near
Campbeltown, ‘‘several hundred feet above sea level”’
(Fifteenth Ann. Rep. of the Fishery Board for Scotland,
Part III., p. 322); and Prof. G. O. Sars says that it
occurs ‘fin brackish pools among shore rocks, consider-
ably above high-water mark,’ and in ‘‘ warm springs of
south Greenland.’ (Amphipoda of Norway, p. 508).

‘“« Pleonexes gammaroides, Sp. Bate.

‘“‘ Several specimens taken at Port Erin by Mr. Chadwick,
in April, 1898, probably in rock pools. Not previously
recorded in the Isle of Man.”

Mr. ArRnoLtD T. Watson, F.L.S., reports to me :—

‘‘ During the past year I have continued the observa-
tions on the habits of Owenia filiformis, to which
reference was made in the last Report, and I am now
working at one or two points which still remain unsettled.
In due course I hope to publish a full account, but mean-

MARINE BIOLOGICAL STATION AT PORT ERIN, 35°

time it may perhaps be of interest to state that, by
establishing a colony of these annelids in my aquarium, I
have been enabled to ascertain their method of reproduc-
tion, and to obtain the larval form. This proves to be
Mitraria, the parentage of which was, I believe, previously
unknown.” }

The Rev. T. 8. za has continued his interesting work
on the preparation of photographic records of the habitats
and appearance of the littoral plants and animals under
natural conditions, either when exposed at low tide or
when actually under water. He has produced a most
beautiful series of plates showing sea-anemones expanded
in shore pools, in some cases catching and swallowing
food. last year Mr. Lea photographed (half-plate size)
a marked area of rock covered with adhering animals,
This year he has taken the same area, on the same scale.
and finds (see fig. 4) that the population has changed
almost entirely. All last year’s limpets are gone, leaving
their scars on the rock; a few of the barnacles seem to
be the same individuals. Many thousands of new animals
have appeared.

Miss L. R. THorRNELY has found on material from
Port Erin the following interesting Polyzoa new to our
district :—Mucronella peachw, var. labiosa, previously
known from Belfast and Guernsey; and Mucronella
abyssicola, on a shell, previously known only from
Shetland.

Dr. G. W. Chaster informs me that from material
dredged in our district by the Fisheries steamer he has
obtained the mollusca Huloma intermedia (alive and very

ol i a el ii -

[voy “SL “At

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

36

Aq sopoyd worq] “yTRUL 19ZVM MOT BAOGE 4o2T 9 Juoge ‘ULIGT Og ‘Lec yourpredg ‘aovj-yoo1 Roa A =“ BLT

i

?

June, 189

June, 1898,

MARINE BIOLOGICAL STATION AT PORT ERIN. 37

fine) and Neolepton obliquatum, the latter a species which
he added to the British fauna from Irish dredgings.

It will be remembered that in our Sixth Annual Report
(December, 1892), I gave an account of the colour varieties
of the little shore prawn, Hippolyte (or Virbwus) varians,
and of their ‘‘ protective’? nature, as found on different
kinds of sea bottoms at Port Erin. A coloured plate (PI.
VI.) showed the green form on the green plant Zostera ;
a red form on the sea-weeds Delesserta and Rhodymema ;
and a dark olive brown form on the bushes of the brown
Alga, Halidrys siliquosa. Four possible explanations of
the facts were given, and it was stated that an experi-
mental enquiry into the matter was in progress.

The following year, in the Seventh Report (1898, p. 35),
the results of the experiments, so far as then obtained,
were given, leading to the conclusion “that the adult
animal can change its colouring very thoroughly, although
not in a very short space of time.’’ A short description
of the conditions of the differently coloured chromato-
phores or pigment cells of the skin during these changes
was given, and their microscopic characters briefly dis-
cussed. It was stated ‘it would be. interesting to
determine whether . . . . the modification of the
chromatophores is due to nerve action and is dependent
upon sight, or is the result of the direct action of light
upon the integument,” but no further experiments bearing
upon this point were then made.

During the following year the experiments were repeated
at Port Erin, with practically the same results. Glass
jars, painted on one side, or covered with coloured papers,
as well as those containing coloured weeds, were used;
and the prawns, while changing colour, were occasionally
examined under the microscope, and some coloured draw-
ings were made of the elaborately branched chromato-

38 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

phores, showing the distribution of the blue, yellow, red,
brown, and chocolate pigment. The whole subject, so far
as then known, was discussed in a lecture at Port Erin,
to the Isle of Man Natural History Society, in the summer
of 1894, and a passing reference to it was made in the
Tenth Annual Report, in connection with other cases of
colour change, such as those of some fishes. There I left
the matter, and I am glad to say that Mr. James Hornell,
in Jersey, and Messrs. F. W. Gamble and F. W. Keeble,
in our own district, have now taken up the investigation,
with the intention of carrying it further.

Messrs. GAMBLE and KEEBLE, both Demonstrators of
Biology in the Owens College, have been making a number
of interesting observations and experiments in regard to the
physiology of the colour changes of Hippolyte and other
prawns during this past summer. Most of their work
has been carried out at the Piel Sea-Fish Hatchery, but
they have in part made use of specimens obtained at
Port Erin. Their results are not yet ready for publication,
but I may remark that the chief novelty in their methods
is the use of water-tight glass chambers, through which
fresh aérated sea-water can constantly pass, and in which
the prawns can be kept under constant observation while
different colours of light are applied. A fuller account of
this method, by Messrs. Gamble and Keeble, will be given
in the forthcoming Annual Report to the Lancashire Sea-
Fisheries Committee.

Professors Boyce and HERDMAN, with the help of Dr.
Koun, have continued during the year their work on
Oysters and Disease, and presented their third, and final,
report upon the subject to Section D of the British
Association at the Bristol meeting in September. Amongst
the conclusions at which they have arrived may be quoted
the following :—

MARINE BIOLOGICAL STATION AT PORT ERIN. 39

“1. There are several distinct kinds of greenness in
oysters. Some of these, such as the green Marennes
oysters and those of some rivers on the Hssex coast, are
healthy; while others, such as some Falmouth oysters
containing copper, and some American oysters re-bedded
on our coast, and which have the pale green leucocytosis
we described in the last report, are not in a healthy state.

“2. Some forms of greenness (e.g., the leucocytosis) are
certainly associated with the presence of a greatly-increased
amount of copper in the oyster, while other forms of
ereenness (¢.g., the Marennes) have no connection with
copper, but depend upon the presence of a special pigment
Marennin, which may be associated with a certain amount
of iron.

“3. We see no reason to think that the iron in the
latter case is taken in through the surface epithelium of
the gills and palps; but regard it, like the rest of the iron
in the body, as a product of ordinary digestion and absorp-
tion in the alimentary canal and liver.

‘““4. We do not find that there is any excessive amount
of iron in the green Marennes oyster compared with the
colourless oyster; nor do the green parts (gills, palps,
&c.) of the Marennes oyster contain either absolutely or
relatively to the colourless parts (mantle, &c.) more iron
than colourless oysters. We therefore conclude that there
is no connection between the green colour of the ‘ Huitres
de Marennes’ and the iron they may contain.

“5. On the other hand, we do find by quantitative
analysis that there is more copper in the green American

-oyster than in the colourless one; and more proportionately

in the greener parts than in those that are less green. We
therefore conclude that their green colour is due to copper.
We also find a greater quantity of iron in these green
American oysters than in the colourless; but this excess

40 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

is, proportionately, considerably less than that of the
copper.

“6. In the Falmouth oysters containing an excessive
amount of copper, we find that much of the copper is
certainly mechanically attached to the surface of the body,
and is in a form insoluble in water, probably as a basic
carbonate. In addition to this, however, the Falmouth
oyster may contain a much larger amount of copper in its
tissues than does the normal colourless oyster. In these
Falmouth oysters the cause of the green colour may be
the same as in the green American oysters.

“7. The Colon group of bacilli is frequently found in
shellfish, as sold in towns, and especially in the oyster ;
but we have no evidence that it occurs in Mollusca living
in pure sea-water. The natural inference that the presence
of the Colon bacillus invariably indicates sewage contamina-
tion must, however, not be considered established without
further investigation.

‘8. The Colon group may be separated into two divisions
—(1) those giving the typical reactions of the Colon bacillus,
and (2) those giving corresponding negative reactions, and
so approaching the typhoid type; but in no case was
an organism giving all the reactions of the B. typhosus
isolated. It ought to be remembered, however, that our
samples of oysters, although of various kinds and from
different sources, were in no case, so far as we are aware,
derived from a bed known to be contaminated or suspected
of typhoid.

“9, Consequently, as the result of our investigations,
and the consideration of much evidence, both from the
oysters growers’ and the public health officers’ point of
view, we beg to recommend :—

‘“‘(a) That the necessary steps should be taken to
induce the oyster trade to remove any possible

MARINE BIOLOGICAL STATION AT PORT ERIN. 41

suspicion of sewage contamination from the beds and
layings from which oysters are supplied to the market.
This could obviously be effected in one of two ways,
either (1) by restrictive legislation and the licensing
of beds only after due inspection by the officials of a
Government department, or (2) by the formation of
an association amongst the oyster-growers and
dealers themselves, which should provide for the due
periodic examination of the grounds, stores and stock,
by independent properly-qualified inspectors. Scien-
tific assistance and advice given by such independent
inspectors would go far to improve the condition of
the oyster beds and layings, to re-assure the public,
and to elevate the oyster industry to the important
position which it deserves to occupy.

““(b) Oysters imported from abroad (Holland,
France, or America) should be consigned to a member
of the ‘ Oyster Association,’ who should be compelled
by the regulations to have his foreign oysters as
carefully inspected and certificated as those from his
home layings. A large proportion of the imported
oysters are, however, deposited in our waters for such
a period before going to market that the fact of their
having originally come from abroad may be ignored.
If this period of quarantine were imposed upon all
foreign oysters a great part of the difficulty as to
inspection and certification would be removed.

‘“‘(¢) The grounds from which mussels, cockles and
periwinkles are gathered should be _ periodically
examined by scientific inspectors in the same manner
as the oyster beds. The duty of providing for this
inspection might well, we should suggest, be assumed
by the various Sea-Fisheries Committees around the
coast.”

42. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Mr. A. R. Jackson, B.Sc., has continued his work on
the Spiders of the neighbourhood of Port Erin, and has
now drawn up a list of species which has been communi-
cated to the Biological Society.

Mr. ANDREW Scott, working chiefly at Piel Island,
has made some interesting observations upon the food of
young fishes. He finds that young Mullet up to an inch
and a little more in length are still feeding entirely upon
Diatoms (chiefly Navicula). At a length of one-and-a-
half inches they feed on both Diatoms and Copepoda
(Tachidius). Young Herring up to three inches long
were found feeding on Mysvs.

Mr. F’. J. CoLE has continued throughout the year his
preparation of material for his joint research with Prof.
Herdman on the process of budding and the formation of
colonies in compound Ascidians. He has visited Port Erin
about four times during 1898 for the purpose of collecting
and preserving old and young colonies of the species
commonly known as Amarouciwwm argus, which, however,
is seen by its anatomical character to belong to the genus
Morcheluum. Myr. Cole and Prof. Herdman have also
kept various kinds of compound Ascidians alive under
observation in the tanks, so as to record the mode of
erowth and the positions of the new buds in the colony.
Before regarding the collection of material as complete, it
has been considered advisable to examine colonies at dif-
ferent times of the year so that no stage of importance might
be omitted. Colonies of Ascidians have therefore been
collected from month to month from Easter, 1897, and
this will be continued until next April. Owing to the
difficulty experienced in satisfactorily staining the pre-
parations, the microscopic part of the work is somewhat
laborious and takes considerable time. Any detailed
statement of results of work would, at the present stage

MARINE BIOLOGICAL STATION AT PORT ERIN. 43

of the enquiry, be premature, and perhaps lacking in
accuracy, but it may be remarked that what is seen in the
first series of sections of the young stages of buds is more
in accord with the conclusions of Ritter and of Lefevre
than of other investigators.

EXTENSION OF THE WORK.

THERE are several matters, in addition to the routine work
of the Station, which call for special attention this year.

One of these is the necessity for further exploration in
the North Atlantic. Attention has repeatedly been drawn
of late years to the importance, both from the purely
scientific and the industrial points of view, of the problems
involved. The Scandinavians (Petterssen, Ekman, Hort,
and others) have succeeded in unravelling some of the
interlacing belts of water from Arctic, Baltic, North Sea,
and Atlantic sources which sweep past their coast, and
affect the movements of migratory fish. It is only by
such work that we can hope to explain rationally the
mysterious movements of the Herring—perhaps the most
important food fish on our coast.

It was formerly supposed* that when the Herrings
left our shores in autumn they retired to the far north, and
next season started from the Arctic regions on their annual
migration, led by one large old fish—the “‘ King of the
Herrings.”” We now believe that breeding and feeding are
the two impulses that govern the movements of a fish.
The Herring comes into shallow water on our coast to
spawn, and when it migrates in search of food from
the Atlantic to the North Sea, or from our West
Coast out into deep water, we have reason to believe
that it is following those minute organisms which

* H.g., see Pennants’ British Zoology, and article Ichthyology in Ency,
Brit. for 1857.

44 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

form the plankton carried along in particular currents
of water, characterised by the temperature, the salinity,
and the microscopic fauna. It is possible by these
characters to recognise the currents, to trace their
variations from year to year, and so to some extent to
determine and predict the movements of the shoals of
fish. We owe this view chiefly to Professor G. O. Sars,
of Christiania. It is, then, to the physical conditions and
the biological contents of the water that the movements
of the Herring are due, and these are matters within the
scope of man’s investigation, but outside his power to
regulate by local bye-laws, imperial legislation, or even
international treaty. This should be recognised by the
fishermen and by Sea-Fishery authorities.

It follows, then, that one of the most important things
the Biologist can do to add to our knowledge of life in the
sea is to make a survey of the microscopic floating and
drifting life of the sea, and its relation on the one hand to
the physical conditions at the time (especially the
temperature and salinity of the water), and on the other
to the food materials found in the stomachs of the fish.

The “‘pump plankton’’ method, which I described fully
before this Society last session, might do much if
systematically worked, but it seems difficult to induce
anyone on a ship, except a naturalist, to undertake such
work, and, moreover, the organisms collected may, unless
great care be taken, suffer so much damage in the process,
as to be difficult of identification. How, then, are we to
sample the plankton of our oceans? The attempt has
been made lately to arrange for the stoppage of a steamer
at certain fixed points in the ocean long enough to permit
of surface and deep-sea gatherings being taken. This
method is difficult to arrange, liable to failure, and very
expensive considering the very small number of observa-

MARINE BIOLOGICAL STATION AT PORT ERIN. 45

tions that could be taken—such as eight in a proposed
traverse from England to the West Indies.

A much more satisfactory plan, from the scientific point
of view, would be to have a series of special cruises, in a
vessel of the type of a fisheries steamer, a steam trawler,
or a steam yacht able to keep at sea, carrying lines of
closely-placed observations across those areas that
influence our British Marine Fauna. I would suggest :—

1. A line from Land’s End to Porto Rico and Jamaica
in the West Indies crossing Rennel’s Current, and the mid-
Atlantic south of the Gulf Stream area, and tracing back
those elements of our floating fauna that we owe to the
5.W. Atlantic drift towards their tropical source.

2. A line from Ushant to Iceland, so as to cross the
entrance to the English Channel and the drift of the
Atlantic organisms on to the west coasts of Ireland and
Scotland, and then across the entrance to the Arctic
Ocean and North Sea.

3. A line from the Shetlands to the southern end of
Greenland, along lat. 60° N., so as to intercept any Arctic
flow which brings down the northern forms found in our
fauna.

The British Islands occupy a central and commanding
position, touching, as Prof. Edward Forbes pointed out
long ago, upon the Arctic and Scandinavian faunas to the
North, and the Germanic, South European, and Atlantic
faunas to the south.

It would also be of considerable scientific interest—
although having no bearing so far as can be seen at
present upon practical fishery questions—to explore more
systematically the seas to the north of Iceland and Norway
as far north as dredging and tow-netting are practicable.
Sir John Murray, it is well known, has recently revived
an old idea that the faunas of the two poles are closely

46 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

similar. In opposition to this ‘‘ bipolar hypothesis,” I
pointed out last session that in the Tunicata at least there
was no special resemblance between the far northern and
far southern species, and that, on the other hand, there
was a marked similarity between a series of forms from
the North Pacific and those from the North Atlantic.
Sir John Murray answers me, in a letter on the subject,
‘“‘T think your remarks quite just. I expect many cases
of north and south distribution will be knocked on the
head by further discoveries, and that many cases of
bipolar distribution not yet evident will be brought to
hight.” Well, we want now to determine how far that is
the case. We both agree that what is needed is further
facts—more investigation.
There is every prospect that the Antarctic Seas will be
fully explored. Several expeditions—Borchgrevink’s and
others—are there or will be soon, still others are in
prospect. The Tunicate fauna of Australian seas is fully
as well-known as that of Europe, the marine fauna round
Kerguelen Island, thanks to the ‘‘ Challenger’’ explora-
tions and Sir John Murray’s writings, is better known
than that of Iceland, whilst of most of the inhabitants of
the seas around Spitzbergen we are practically ignorant.
Moreover, Nansen has opened up a new problem in
northern marine exploration. He has shown that a deep
sea basin occupies a part at least of the polar area. Where
does that basin begin, and how far does it extend? How
does it end towards Norway, Iceland, and Greenland ?
The sea immediately to the north and west of Norway is
shallow, running out gradually to a depth of about 200
fathoms, and then, according to Mohn, descending steeply
to the bed of the deeper ocean, which reaches a depth of
2000 fathoms or more. We do not yet know the limits
nor the inhabitants of that deep basin. In advocating

MARINE BIOLOGICAL STATION AT PORT ERIN. AT

Antarctic expeditions we must not forget how much still
remains to be done within the Arctic circle.

All this may seem to have little connection with our
L.M.B.C. work, but it is the natural extension and out-
come of what we have been doing. As the problems
develop we must widen our area. We commenced twelve
years ago with Hilbre Island and Liverpool Bay—our
work has now for several years extended all over the Irish
Sea. This means practically the western fauna of Britain
influenced by the Atlantic drift from the south-west and
the Arctic currents from the north. Whether the relation
of our north-western Huropean fauna to that of north-
western America depends upon a common circum-polar
fauna as I have suggested, and whether our thorough
comprehension of the Arctic fauna is bound up, as Murray
thinks, with Antarctic explorations, are questions still to
be answered; but this much is clear, that one enquiry in
the distribution of animals leads to another, and the
different faunas are like links in a chain or strands in a
net—the mesh-work of life extending over the globe—
just what we should expect from the consideration that
all living things spread from a centre, that multiplication,
strugele for existence, migration, survival of the fittest,
and varying degrees of isolation have produced the dif-
ference we now find between the present inhabitants of
the different regions.

I should like, then, to push our L.M.B.C. investigations
further afield—out into the North Atlantic, across to
the West Indies, up into the Northern Seas. And all
that is wanting is a fund to meet the necessary expenses.
We can find the scientific men willing to give the time
and do the work. What we want is a yacht-owner willing
to use his vessel, fitted up with the necessary equipment,

48 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

for scientific exploration of an interesting and important
nature, such as the Prince of Monaco is now doing.

There are two practices in American universities which
excite the envy of professors in this country. One is the
‘‘sabbatical year’’—the one year in every seven given for
purposes of travel, study, and investigation. The other
is the frequent endowment of an expedition—or equipment
of an exploring party—by an individual man or woman
who is interested in the subject, and can give a special
fund for such a purpose. The Columbia University in
New York, the Johns Hopkins University in Baltimore,
Yale College in Newhaven, and Harvard at Cambridge,
have all benefitted immensely in the past by such explor-
ing expeditions. Nearly every year of late has seen one
or more of such, due to private generosity, in the field;
and the work they have done has both added to general
scientific knowledge, and has also enriched with collections
the laboratories and museums of the college to which the
expedition belonged.

It may be that the reason why this excellent system of
exploring parties from the universities has attained such
slight development in our country is that our professors
are not so enterprising in making known the wants of
science—not so importunate in their demands upon the
community in which they live—as their transatlantic
brethren. It may be that some think that in this long-
settled country there is nothing left to explore—no
greater mistake could be made. I am sure that the
geologist and the archeologist could point to innumerable
problems still to be attacked on land, while our seas are
fay more vast, and comparatively far less known than
the shores. I am sure that many marine zoologists
could be usefully employed during their vacation for
the next ten years in exploring such regions as I have

MARINE BIOLOGICAL STATION AT PORT ERIN. 49

indicated above. While, if we pass from questions of dis-
tribution to those of structure and life-histories, even the
best-known regions afford abundant opportunity for
research to all who enter the field.

Marine Biology has this peculiar advantage, that invok-
ing as it does the aid of several neighbouring sciences,
such as Geology, Geography, and Physiology, it is able in
its turn to throw light upon these subjects, and it bristles
with problems of that interesting and intricate nature that
characterises the borderland between two or more sciences.

SomE REcENT WORK.

As a contribution to the borderland between our subject
and Geology, Mr. Lomas and I have, during the past
year, made a careful examination of all our dredged
deposits from the floor of the Irish Sea, and our results
have appeared in the last volume of ‘‘ Proceedings”’ of
the Liverpool Geological Society. Amongst our results
we may note that we have shown how the material now
covering the floor of the Irish Sea have been the result (1) of
the denudation of the coasts, (2) of the redistribution of
the older deposits under the sea, (3) of vital agencies—
the remains of animals and plants living in the sea. We
give a revised classification of such deposits, and a detailed
description of all the samples we have obtained.

As a problem on the boundary between Biology and
Geography (viz., the currents of the sea), and as having,
at the same time, an important practical application to
the sea-fisheries, may be cited our investigation of the
circulation of water in the Irish Sea by means of “ drift-
bottles.”” Our records of those bottles that have been
found and returned to us show pretty conclusively that
there is, in addition to both north and south in-flowing
and out-going tidal currents, a slow drift to the north,

50 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

which would carry, for example, floating objects, such as
fish eggs and embryos, gradually further and further to
the north. So that, from the spawning grounds south of
the Isle of Man, they may be carried towards and along
the Lancashire coast and from say the ‘‘ Hole”’ and other
parts of the central area, to Cumberland and the Solway
Firth.

As another research intimately bound up with the local
fisheries, I would allude to our plankton investigations.
During the past year surface gatherings have been sent
from various localities with more or less regularity. The
result of the examination of these, on the whole, confirms
very well-the general conclusions we drew in the last
Report as to the succession of forms throughout the year.

We find very considerable difference between gather-
ings from different localities; for example, whereas the
tow-nettings from Port Erin are clear and clean and
support an abundant assemblage of minute animals,
those taken about the same time off Peil, in the Barrow
Channel, contain much vegetable débris, mud, &c., due to
the influence of fresh-water and the washings of the land.
The true open sea at Port Erin and the absence of any
body of fresh-water, and of any mud flats, ensure the
presence of a much larger number of Copepoda, Dino-
flagellata, and other characteristic pelagic organisms than
are elsewhere. We have also noticed that the various
constituents of the surface life—both larval and adult—
appear earlier at Port Erin than on the Lancashire coast.

Tur Manx SEA-FISHERIES COMMISSION.

The report which recently appeared containing the
recommendations of the Manx Industries Commission, so
far as they relate to sea-fisheries, includes an important
paragraph in which it is suggested that the Hatchery

MARINE BIOLOGICAL STATION AT PORT ERIN. 51

which is contemplated be established in connection with
our Biological Station at Port Erin. The possibility of
some such arrangement being proposed, caused the
Committee to consider carefully the ground in the
neighbourhood of the Station with the view of determining
how far they could offer accommodation for economic
work, and what additional building and plant would be
required before they could undertake sea-fish hatching on
anything beyond an experimental scale. The chief
requirements for such a purpose are as follows :—

1. An extension of the Aquarium house, to hold the
hatching boxes.

2. A small boat jetty.

3. A concrete pond on the shore.

4. A circulation of sea-water.

All of these are merely a matter of expense. They can
readily be provided if a grant be given for the purpose.
The hatching-house will cost about £80, the jetty £70,
and the pond £80; say, including the fittings of the
hatchery, £250 in all. | :

As to a constant flow, or circulation, of sea-water, there
is no gas supply in Port Erin, and if worked by a steam
engine a very considerable expenditure and the services of
an engineer would be required. A much more economical
plan, and one that would probably be quite sufficient for
the wants of the institution, would be to pump from the
sea-well to the uppermost tank by means of a small wind-
mill, as is done in some of the Biological Stations abroad.
This would be comparatively inexpensive, and would not
require material addition to the service of the institution ;
our present Curator with the assistance of a strong lad,
and possibly a second fisher lad during the hatching
season, would be able to undertake the work.

52 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

L.M.B.C. PuBLIcaTions.

During the past year the following papers dealing with
L.M.B.C. work have been published :—

1. The Eleventh Annual Report (Trans. Biol. Soc.,
Vol GXGiS Oil):

2. The Sea-Fisheries Laboratory Report for 1897
(Trans. Biol. Soc., vol. XII., p. 176).

38. Note on a Tetramerous Specimen of EHchinus
esculentus. By H. C. Chadwick (Trans. Biol. Soc.,
vol. XIT., p. 288).

4. Actinological Studies, I.—The Mesenteries and Ciso-
phageal Grooves of Actinia equina. By J. A. Clubb,
M.Sc. (Trans. Biol. Soc., vol. XII., p. 300).

5. Report on Oysters and Disease, by Prof. Boyce and
Prof. Herdman (Brit. Assoc. Rep.), with an Appendix on
Iron and Copper in Oysters, by Dr. Kohn.

6. The Deposits on the Floor of the Irish Sea. By
Prof. Herdman and Mr. Lomas (Proc. Liverpool Geol.
Soc. vole p. 205):

It is hoped that volume V. of the ‘‘ Fauna and Flora,”’
containing reprints of these and all other papers on our
local work published since the appearance of vol. IV. Gn
1895) will be ready to issue during the present winter.

I am inclined to think that the time has now arrived
when it may be the duty of the Committee to issue a new
form of publication. Our work hitherto, during the last
12 years, has been largely faunistic and speciographic.
The work of necessity must be so at first when opening
up a new district. Some of our workers have published
papers on morphological pcints, or observations on life
histories and habits; but the majority of the papers in
our volumes on the Fauna and Flora of Liverpool Bay
have been, as was intended, occupied with the names and
characteristics and distribution of the many different

%
q

MARINE BIOLOGICAL STATION AT PORT ERIN. 53

kinds of plants and animals in our marine district. And
this Faunistic work will still go on. It is far from
finished, and I hope we shall still add greatly to our
records of the Fauna and Flora; but in addition it might
be useful to produce a series of

L.M.B.C. MEmoIrrs.

What I now propose is that each of our specialists
should carefully prepare a full account, illustrated by all
the necessary figures, of one or two common but
important animals belonging to the group upon which he
is an authority; and that these detailed and fully
illustrated accounts should be issued as a series of
L.M.B.C. Memoirs — obtainable at first separately,
memoir by memoir as they appear, and then later bound
up, say 5 or 6 together, in convenient volumes. I believe
that such a series of special studies, written by those who
are thoroughly familiar with the forms of which they
treat, will be of great value to students in our laboratories
and in Biological Stations, and will be welcomed by
many working at marine biology. It is proposed that the
forms selected should, as far as possible, be common
L.M.B.C. (irish Sea) animals of which no adequate
account already exists in any text book. Probably all
the members of our band of specialists will be willing to
help in this work. The following have already promised
their services, and will probably treat of the species
placed opposite their names:—

Professor Harvey Gibson Zostera marina.
Mr. C. E. Jones ... ... Haldrys siliquosa.

Professor Weiss ... ... Typical Diatoms (Biddul-
phia, Chaetoceros, Schiz-
onema, and Liemophora).

54 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Dr. O. V. Darbishire ... A Red Sea-weed.

Mri. JévCole ia. ... The Plaice.

Professor Herdman ... Ascidia and Botrylloides.
Mr. W. E. Hoyle .. A Cuttle-fish (Hledone).
Mr. J. Johnstone... ... The Cockle.

Mr. Andrew Scott .... AnOstracodandan Epizoon.
Mr. I.C. Thompson... Calanus firmarchicus.
Mr. H. C. Chadwick ... Echinus, the Sea-urchin.
Mir Sete Club tx: ... An Actinian.

Miss L. R. Thornely .... A Polyzoon Colony.
DrEtaniisch mine ... -A Calcareous Sponge.

Now, all these memoirs will require to be carefully
illustrated with well-drawn plates, and fortunately the
Committee have at their disposal a small annual sum of
money (the British Association—1896—fund) which it
was proposed from the first should be devoted to just such
a purpose as this. That, however, leaves the printing of
the memoirs still unprovided for, and with the Curator’s
salary and other expenses at Port Erin to meet, it will be
quite impossible to spare anything for general funds. A
special subscription will have to be raised to defray the
expenses of the L.M.B.C. Memoirs.

I should also hke to point out that papers on special
points in the Zoology of Sea-Animals are being sent by
our workers to London Scientific Societies, because we
have not the funds to publish them satisfactorily in
Liverpool. Mr. Andrew Scott has just finished a beauti-
ful series of water-colour drawings of fish eggs, embryos
and larve, from his investigations during the year at
our Lancashire Fish Hatchery. They will be. ex-
pensive to reproduce and publish, but it will be very
discreditable to Committee, and City, and County, if
we allow this excellent local work to be issued under

MARINE BIOLOGICAL STATION AT PORT ERIN. ND

the auspices of some society in London or Edinburgh.
The fact is—and a very important and gratifying fact it
must be to our subscribers and supporters who
have helped us from year to year—that we have
now in connection with our Committee an exceed-
ingly active School of Marine Biology, every worker
in which is engaged in some piece of original
research. Liverpool is the right and natural place for
a School of Marine Biology, and I hope that Liverpool
will consider that it is creditable to the city that such
local researches should be published by a Liverpool
Society. An addition of about £100 a year to our
funds is necessary in order to enable us to do justice
to the work now being produced by our colleagues and
students. :

The Biological exploration of Liverpool Bay has all
along been intimately connected with University College,
and it is to be hoped that whenthe time comes—and it
ought not to be long delayed—for building new biological
laboratories, accommodation will be provided within the
department for our School of Marine Biology. A museum
devoted to exhibiting the products of the local seas, and a
laboratory fitted for conducting researches upon marine
life, must surely be constituent parts of the University of
a great sea-port.

56 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

APPENDIX A.

THE LIVERPOOL MARINE BIOLOGY
COMMITTEE (1898).

R. D. DARBISHIRE, Esq., B.A., F.G.S., Manchester.

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

His ExcELLENCY LoRD HENNIKER, Governor of the Isle
of Man.

Pror. W. A. HerpMay, D.Sc., F.R.S., F.L.8., Liverpool,
Chairman of the L.M.B.C., and Hon. Director of
the Biological Station.

W. EK. Hoye, Esq., M.A., Manchester.

A. LEICESTER, Esq., formerly of Liverpool.

Sir JAMES Pooue, J.P., Liverpool.

Dr. Isaac Roserts, F.R.S., formerly of Liverpool.

I. C. THompson, Esq., F..8., Liverpool, Hon. Treasurer.

A. O. WALKER, Esq., F.U.5., J.P., Colwyn Bay.

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

I.—The Oxpsect 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.

MARINE BIOLOGICAL STATION AT PORT ERIN. 57

II.—The CommiTT#E shall consist of not more than 12
and not less than 10 members, of whom 3 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.

I1I.—During the year the Arrairs of the Committee
shall be conducted by an Hon. Director, who shall be
Chairman of the Committee, and an Hon. TREASURER,
both of whom shall be appointed at the Annual Meeting
and shall be eligible for re-election.

IV.—Any VACANCIES 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 Expensss 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 BroLocicaL Srarion shall be used primarily
for the Exploring work of the Committee, and the
SPECIMENS collected shall, so far as is necessary, be
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.

58 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

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. I. C. Thompson, F.L.S.).

Ii.—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 or Laboratory
Assistant, 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, boat, collecting apparatus,
&c., are to be employed.

Il1I.—The Resident Curator will be ready at all
reasonable 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, to see the Aquarium and the Station, so
long as it is found not to interfere with the scientific
work. Occasional lectures are given by members of the
Committee.

V.—Those 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 four 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 Colleges and Museums, may become

MARINE BIOLOGICAL STATION AT PORT ERIN. 59

subscribers in order that a work place may be at the
disposal of their 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.—EHach worker* is entitled to a work place opposite
a window in the Laboratory, and may make use of the
microscopes, reagents, 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.

VIL.—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 he
taken away from the Laboratory. Workers desirous of
making, preserving, and 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
to 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
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

* Workers at the Station can always find comfortable and convenient

quarters at the closely adjacent Bellevue Hotel; but lodgings can readily be
had by those who prefer them.

60 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

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 duplicate 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 communi- -
cations relating to finance, should be sent to the Hon.
Treasurer, Mr. I. C. Thompson, F.L.8., 53, Croxteth
Road, Liverpool. Applications for permission to work at
the Station, or for specimens, or any communication in
regard to the scientific work should be made to Professor
Herdiman, F'.R.8., University College, Liverpool. }

LANCASTER e

IRELAND pig. ae Ein a. ot . we

Diagrammatic Section across the Irish Sea..

MARINE BIOLOGICAL STATION AT’ PORT ERIN. 61

APPENDIX B.

HON. TREASURER’S STATEMENT.

From the balance sheet appended it will be seen that
the year ends with a small debit balance, in spite of the
fact that all due economy has been observed, consistent
with efficiency.

The British Association (1896) fund has, to a consider-
able extent, enabled the Committee to meet the necessary
extra expense involved in securing the services of a com-
petent Resident Curator at the Port Erin Station, which
has already proved of great benefit to workers.

Increased workroom accommodation has also been
added at the Station during the year, at a cost of about
£14.

As pointed out by the Director in his Report (p. 55), an
additional income of about £100 per annum could be
most advantageously utilized in publishing well-illustrated
papers and memoirs, embodying the results of local bio-
logical investigations such as, for want of funds, have
unfortunately, in some cases, been sent, in the past, to
other societies outside Liverpool.

The Hon. Treasurer, on behalf of the Committee, will
be glad to receive the names of new subscribers, with the
view of remedying this deficiency, and of further adding
very materially to the already excellent work achieved
under the auspices of the L.M.B.C. since its foundation,
twelve years ago.

62 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

SUBSCRIPTIONS and DONATIONS.

Subscriptions. Donations.
sv fides d

Ayre, John W., Ripponden, Halifax te
Bateson, Alfred, Harrop-road Bowdon... 1 1 0
Beaumont, W. I., Cambridge fama Pats)
Bickerton, Dr., 88, Rodney-street ale ne
Bickersteth, Dr., 2, Rodney-st. ... sos 2
Brown, Prof. J. Campbell, Univ. Coll.... 1-1 0
Browne, Edward T., B.A., 141, Uxbridge-

road, Shepherd’s Bush, London Teen
Brunner, Sir J. T., Bart., M.P., Druids

Cross ... 5 0 0
Boyce, Prof., iti oeeat Gpllase Wee ae
Caton, Dr., 86, Rodney-street — iL
Clague, Dr., Castletown, Isle of Man Tt ee
Clague, Thomas, Bellevue Hotel, Port Erin 1 1 0
Cole, F'. J., (research table) wie “le ealema
Coombe, John N., 4, Paradise-sq., Sheffield Lt te
Comber,Thomas,J.P., Leighton, Parkgate 1 1 0O
Crellin, John C., J.P., Ballachurry, An-

dreas, Isle of Man 3 0 10) 36
Crossland, Cyril, Clare College, ae 1 Gee
Gair, H. W., Smithdown-rd., Wavertree 2 2 0O
Gamble,Col.C.B.,Windlehurst,St.Helens 2 0 O
Gamble, F.W.,OwensCollege, Manchester 1 1 O
Gaskell, Frank, Woolton Wood... 2) Sl Sia
Gaskell, Holbrook, J.P., Woolton Wood 1 1 O
Gibson, Prof. R. J. Harvey, 5, Adelaide-

terrace, Waterloo liga
Glynn, Dr., 62, Rodney-street 2° 2 Re
Gotch, Prof., Museum, Oxford ... pe Des

Forward ...£382 13 6 1 0

MARINE BIOLOGICAL STATION AT PORT ERIN. 63
Subscriptions. Donations.
fae Se Ue. en St Ge
Homgarde.. s2 lo 6 tf 1 0
Halls, W. J., 35, Lord-street Ld,..0 —
Hanitsch, Dr., Museum, Singapore EE La) —
Henderson, W.G., Liverpool Union Bank 1 1 O —
Herdman, Prof., University College ee (0) =
Hewitt, David B., J.P., Northwich LCE | 0 —
Holland, Walter, Mossley Hill-road Ze 2 0 —
Holt, Alfred, Crofton, Aigburth ... i MA 0) ——
Holt, Mrs. George, Sudley, Mossley Hill 1 0 O =
Hoyle, W. H., Museum, Owens College,
Manchester He 50 —
Isle of Man Natural echo afd Cote
quarian Society Death eae) a
Jarmay, Gustav, Hartford aS Sue 0) —
Jones, C.W.,J.P., Field House, oe LevO, KO —
Kermode, P. M. O. Hill-side, Ramsey... 1 1 O —
Lea, Rev. T. Simcox, 3, Wellington-fields 1 1 0 —
Leicester, Alfred, Buckhurst Farm, Eden-
bridge, Kent ... be yds, £0 —
Macfie, Robert, Airds os L020 —
Meade-King, H. W.,J.P., Sandfield Baek LISTE eG) ——
Meade-King, R. R., 4, Oldhall- street 010 O —
Melly, W. R., 90, Chatham-street LPL. 40 —
Monks, F. W., Brooklands, Warrington 1 1 0O ==
Mundy, Randal, Manchester OL 106 —
Muspratt, E. K., Seaforth Hall ... DNS) —
Newton, John, M.R.C.S., 44, Rodney-st. 010 6 ==
O’Kell, Robert, B.A., Sutton, Douglas... 1 1 O =
Paterson, Prof., University College ty LO —
Poole, Sir James, Tower Buildings ALY) —
Rathbone, Mrs. Theo., Backwood, Neston 1 1 0O —
Rathbone, Miss May, Backwood, Neston 1 1 0O —
Forward ... £68 9 0 11 0

64 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Subscriptions. Donations.

£ 8, do hosasaes
Forward -... 68._9° 0) fee
Rathbone, W., Greenbank, Allerton 220.0. —
Roberts, Isaac, F.R.S., Crowborough feel ee 0 —
Shackleford, Rev. L. J., Clitheroe 010 0 —
Simpson,J. Hope, Annandale, Aigburth-dr 1 1 0 —
Smith, A. T., junr., 24, King-street T. SRG —
Talbot, Rev. T. U., 4, Osborne-terrace,

Douglas, Isle of Man 1050 —
Thompson, Isaac C., 53, Ghose noua piers (i) —-
Thornely, James (the late), Baycliff,

Woolton iL i 6 —
Thornely, The Misses, Baycliff, Woolton 1 felt —
Toll. d. Mop Korby tPark, Karbye 2 ss oi ae —
Torrance, Gilbert, North Quay, Douglas 1 1. 0 —
Walker, A. O., Nant-y-glyn, Colwyn Bay 3 3 O —
Walker, Horace, South Lodge, Princes-pk. 1 1 0 =
Walters, Rev. Frank, B.A., King William

College, Isle of Man.. I yee —
Watson, A. T., Tapton- OF saci Sheffield 1 Jaleo —
Weiss, Prof. F. E.,Owen’s College, Man’tr. 1 1 0O a
Westminster, Duke of, Katon Hall 5° 01 C —
Wiglesworth, Dr., Rainhill > via): es ee) —
Yates, Harry, 75, Shude-hill, Manchester 1 40" 30 —

£95, 18 0 2 See

SUBSCRIPTIONS FOR THE Hire or ‘‘ WorK-TABLES,’’ OCCUPIED

BY COLLEGES, &c.

Owens College, Manchester
University College, Liverpool

£10" 090
10: One

£20 0 0

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Om bee sae <

"unae “ALINS “LV

*79a4409 PUNO, pun pajripny

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—! pun JUSUIASOAU] JUSUIMOpU

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66

[WORK FROM THE PORT ERIN BIOLOGICAL STATION. ]

LIST of the ARANEIDA of PORT ERIN and
DISTRICT.

By A. R. Jackson, B.Sc. (Vict.),
Victoria University Scholar in Zoology.

WHILE occupying the University College work-place in
the Port Erin Biological Station during the session 1897-
98, in addition to my study of marine life, I paid some
attention to collecting the spiders of the neighbourhood.

Prof. Herdman suggested that I should follow this up,
on a further visit to the station, with the object of com-
pleting a list of the Araneida found at Port Erin; and I
am now, as the result of my observations, able to record
the following 57 species. They were all found within a
radius of 1 mile of the Biological Station, except Amauro-
bius ferox and Hpeira quadrata.

ARANEIDA.
I. DyYSDERIDZ.

Dysdera crocota, rare.
Harpactis hombergit, fairly common.
Segestria senoculata, very common.

II. DRASSIDz.

Micaria pulicaria, fairly common.

Prosthesima latreillet, fairly common.

P. pusilla, rare, only two specimens were taken.
Drassus lapidosus, very common.

D. cupreus, very common.

D. troglodytes, fairly common.

Clubiona terrestris, fairly common.

C, reclusa, fairly common,

ARANEIDA OF PORT ERIN AND DISTRICT. 67

III. AGELENIDZ.

Tegenaria derham, very common.
Textriz denticulata, fairly common.

IV. DIctTynipz.

Amaurobwus fenestralis, very common.
A. similis, very common.
A. ferox, very local, found in Ballasalla quarries.

V. THERIDIDA.

Theridion sisyphwm, common, and only taken in
Colby glen.

Phyllonethis lineata, very common.

Asagena phalerata, one specimen, on the moors.

Ero furcata, only cocoons found.

Pholcomma gibbum, one specimen.

Linyphia triangularis, very common.

L. pusilla, one specimen, on the cliffs.

L. lineata, fairly common.

L. clathrata, fairly common.

Leptyphantes leprosus, very common.

L. minutus, very common.

L. blackwallu, fairly common.

LL. tenuis, fairly common.

Bathyphantes concolor, very common.

B. variegatus, fairly common.

Neriene rubens, fairly common.

N. bipunctata, fairly common.

Hrigone atra, very common.

EL. dentipalpis, very common.

Pedanostethus lividus, fairly common, in gorse
bushes.

Tiso vagans, rare.

68 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Ceratinella breve, fairly rare.
Peponocranium ludicrum, very common.
Pachygnatha degeeri, common.

VI. EHPErrriv&.

NEE:

Vai

Meta menardw, in Bradda mines, very local.

M. meriane, very common.

M. segmentata, very common.

Zilla xz-notatum, very common.

Eipeira umbratica, very common and local, on
cliffs.

i. diademata, very common, local.

EH. quadrata, very common, and found at Fox-
dale.

T'HOMISIDZ.

Xysticus cristatus, very common everywhere.

LYcosIpD&.

Lycosa ruricola, very common.
L. terricola, very common.

L. pulverulenta, very common.
Pardosa nigriceps, fairly common.
P. pullata, very common.

P. amentata, very common.

P. palustris, very common.

IX. ATTIDZ.

Hpiblemium scenicum, rare, two specimens found
on laboratory wall.
Heliophanus cupreus, fairly common.

69

Report on the INVESTIGATIONS carried on in 1898 in
connection with the LANCASHIRE SEA-FISHERIES
LABORATORY at University College, Liverpool, and
the SHA-FIsH HatcHERY 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, Resident Fisheries Assistant at Piel, and Mr.
JAMES JOHNSTONE, Fisheries Assistant at Liverpool.

With Plates I. and II.
[Read March 17th, 1899.]

CONTENTS.
Introduction - 2 : = 2 2 =) 269
Fish Hatching Work at Piel - - - ee : 80
Observations on Leptocephalus - - - - : eso
Observations on Habits and Food of Young Fishes : =a 90
The Plankton Work - : i 5 2
Experiments with Weighted Drift Bottles - - - - 98
The Spawning of the Mussel - - - : - : = 104
On Sea-Fish Hatching - = : : 5 122
Oysters and Disease - - = > Ss Fs £ EO
Iron and Copper in Oysters - - : - = = - 135
Mussel Beds and Mud Banks - - a GIG)
Report from Messrs. Keeble and enablers on Work done at Piel 149
Appendix :—Laboratory Regulations at Piel - . - - 154
INTRODUCTION.

TuIs is the first complete year of Mr. Scott’s work at the
Piel Hatchery and of Mr. Johnstone’s work at the
Liverpool Laboratory, and a glance merely at the above
table of contents will show the extent and variety of the
investigations that have been undertaken.

Mr. Scott’s work has been in the main economic—the
hatching of eges of marketable fishes and the improve-
ment of apparatus and fittings so as to carry on such
work more efficiently in the future. He hasalso, however,
examined many tow-net gatherings and other samples of
the organisms in the waters and on the shores of our
district, and has carried out a further series of experi-

70 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

ments with large ‘‘drift’’ bottles. These, and also his
observations upon young stages of the Eel, are given
below in separate sections. In addition to Mr. Scott’s
own work at Piel we have a first report from Messrs.
Gamble and Keeble, Demonstrators of Biology at the
Owens College, Manchester, upon the interesting work
that they were conducting at Piel during a considerable
portion of last summer.

In one of his letters to me at the beginning (June) of
the research Mr. Gamble says: ‘‘ We hope by the end
of summer to have some sound experimental evidence of
the nature of colour change in Virbiws and possibly in
Mysis as well. . . . Our apparatus consists partly of
an arrangement for maintaining a constant stream of ~
water through a series of observation vessels and partly
of an arrangement by which air is sucked through a
second set of vessels. Thus we can determine whether
change of air or change of water is the most conducive to
health. Then in addition to a dark box we have a series
of ‘light-filters’ by means of which we can obtain the
influence of monochromatic hght on Virbzus both in the
above-mentioned vessels and under the microscope.

For exact physiological work, the sea-water
and gas laid on in the Laboratory at Piel are an immense
advantage—to say nothing of the excellent accommoda-
tion, and from what we managed to do a few weeks ago
both Keeble and I hope that before the end of September
we may have settled some important questions in
connection with the colour-reactions of Crustacea to
various stimuli.” For further particulars I refer to the
authors’ report given below (p. 149). ,

This work has, at present, no obvious connection with
economic fishery problems, but it is impossible to foretell,
in these days of rapid advance in discovery, and in the

_SEA-FISHERIES LABORATORY. Vals

application of science to industries, how soon the results
of what now seems an investigation In pure science may
turn out to have some important practical bearing. For
my part I am of opinion that all knowledge of animal life
In our seas is of importance, and will help us to
understand the life and ways of fishes.

That a detailed knowledge of the sea and its contents
(however minute) must be the basis of fishery practice
and regulation is recognised in the following extract from
the Memorial in favour of International Oceanographic
Exploration sent last April by the Swedish Government to
our Foreign Office :—‘“‘ All fishing in the North Atlantic,
aud especially the presence of the migratory fishes,
depends upon the great currents in the upper layers of the
sea, and the variation of the presence in these layers of
the food required by the fishes, viz., ‘ Plankton’ or organ-
isms of animal or vegetable origin floating in the water.
A knowledge of these currents and of the quality and_
quantity of food they contain is necessary in order to deter-
mine the legislation required for the creation of a rational
organisation of the fisheries.” Then, again, the migrations
of the Cod towards the Lofoten banks and fjords, and of the
winter Herring into the Skagerack, seem, according to
Otto Pettersson, to be regulated by the impact of cold
Arctic and west Atlantic water in winter driving the fishes
to those parts of the sea where the conditions are less
unfavourable.

It is considerations such as these that lead naturalists
to urge that fishery observations and investigations must
not be restricted by any territorial or administrative
boundaries, but should be extended to off-shore waters,
and even the high seas, so as to follow up and unravel the
factors that contribute to the distribution of our coast
fisheries,

72 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Mr. Ascroft has also stayed some time at the Piel
Hatchery, and we include in this Report a short account
from him of his observations on the separation of mud
from sea-water by shell-fish and on the growth of mud
banks upon beds of shell-fish, especially Mussels.

It may be well to note here that the well-equipped
biological Laboratory with tank-house at Piel is now
open, upon certain conditions, to duly qualified investiga-
tors. In addition to the Laboratory accommodation, the
building provides dining-room, writing-room or library,
and about eight bed-rooms for workers. No charge is
made for residence, and meals are provided at a fixed and
reasonable rate. The regulations for workers, as approved
by the Committee, are appended to this Report.

During the course of his work in hatching Cod, Plaice,
and other fish at Piel last spring, Mr. Scott made a number
of coloured drawings of the various embryonic and larval |
stages of our common fishes. We hope that these may be
published on a future occasion, when the series is more
complete.

Having succeeded so far, and shown that the work can
be conducted with the sea-water pumped up at Piel, Mr.
Scott is naturally anxious to be supplied with a pond in
which to keep spawning fish, and with a proper outfit of
hatching boxes, so as to be able to carry on operations on
a much larger scale. As Capt. Dannevig in Norway and
the Scottish Fishery Board (lately at Dunbar, and for
the future at Aberdeen) have adopted a special form of
hatching apparatus, in which the little cubical boxes con-
taining the developing eggs are rocked up and down
constantly in the water of a larger tank, with, so far as
we can ascertain, very good results, it certainly seems
desirable that we should try some of these ‘‘ Dannevig
rocking boxes”’ at Piel, for comparison with the simple

SEA-FISHERIES LABORATORY. 73

tanks with constant flow of water, which we used last
year. Five sets of rocking boxes have, therefore, been
ordered from Norway, where they are made under the
direction of Capt. Dannevig, and are expected to arrive at
Piel in a few days. They wiil be set up along the east
wall of the tank-house, and the plain tanks, which will
also be in use this coming season, are now being moved into
the adjoining portion of the verandah, which is being
enclosed for the purpose so as to form an extension of the
tank-house.*

The provision of a satisfactory spawning pond, in which
the parent fish can be kept in considerable numbers until
they produce their eggs, is a more difficult matter. The
Scientific Sub-Committee have had the matter under
consideration at several meetings, and I have gone care-
fully into the question of alternative sites, on the ground,
with the Chairman, the Superintendent, Mr. Scott, and
the Engineer to the Railway Co. (our landlords at Piel).
The difficulty is to get a site which is sufficiently near to
the hatchery, sufficiently protected from heavy seas,
which can be excavated to a sufficient depth, and which
will enable the pond to be constructed at a reasonable
cost. A tidal pond, such as that now being constructed
at Bay of Nigg, Aberdeen, by the Fishery Board for
Scotland, has certain obvious advantages, the chief of
which is that no pumping is required, but it 1s open to
the objection that it must of necessity (at Piel) be on the
shore, and, therefore, exposed to the seas.

We are trusting for one year more to such supplies of
spawn as can be obtained by the steamer or from the
trawlers, but if these supplies are as poor as they were
last year, we shall require in early summer, at the latest,

* Since the above was written the hatching boxes have arrived, and the

necessary changes in the accommodation have been carried out.

74 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

to decide upon the position and nature of the spawning
pond, in order that it may be completed in time for the
following season.

Mr. Johnstone’s work at Liverpool has been partly
the investigation in the Laboratory of any specimens
that were sent to us, partly assisting me in the
work, and partly (with Mr. Tom Mercer) looking after
the Fisheries Museum, and the Circulating Fisheries
Exhibition now at Preston. A good deal of Mr.
Johnstone’s time in the Laboratory has been taken
up with the examination of Mussels and other shell-fish,
at all times of the year, and from various parts of the
district. The evidence as to the spawning habits of the
Mussel obtained from the microscopic characters of the
reproductive organs is rather puzzling, but it now seems
most probable that the Mussel commences to produce
mature reproductive elements in the middle of winter,*
and continues to emit eggs and sperms in small quantities
for the first six months of the year (probably in increasing
amount after April), during which time large eggs are
always to be found in the ovary; and then, in the middle
of summer, produces rapidly a much greater number of ova,
so as to clear out the contents of the ovary, which suddenly
(about end of July) undergoes a great change, easily
recognisable under the microscope as the ‘‘ spent”’ con-
dition. It then, after a brief interval, proceeds to develop
fresh ovarian tubules, and load up rapidly with young ova,
which develop during the autumn and early winter in
time to be mature at the end of the year. The full details
of these changes in condition will be found in Mr. John-

* We have obtained Mussels with completely formed active spermatozoa
in the middle of December. Mr. Ascroft has found the free-swimming larva
at Lytham in April and May, and the first ‘‘strikes” of young Mussels
are frequently found here upon Alge in June.

SEA-FISHERIES LABORATORY. ! 75

stone’s section of the Report at p. 104, and the various
stages are illustrated by two plates. I have also started
Mr. Johnstone on an investigation of the structure of our
edible Cockle, and hope in next year’s Report to publish
his full detailed account of that animal.

Our travelling “ Fisheries Exhibition” is working its
way through the more important towns of the county.
At the beginning of March, 1898, after a very successful
period at Liverpool, when it was repeatedly visited by
many interested in the subject, the exhibition was trans-
ferred to the Royal Museum, Salford, where it remained
till the end of October. In the meantime, a circular was
issued from the County Offices, Preston, stating that this
Fisheries Exhibition might be obtained on loan, on certain
terms to meet the expenses of packing, unpacking, re-
arranging, and carriage. Applications were received from
the Museums of Preston, Warrington, Bolton, and St.
Helens; and early in November the exhibition was trans-
ferred to the Harris Free Museum at Preston, where
it will remain for some months. The transference of the
cases and collections (which have to be very carefully
packed and unpacked) from one institution to another
occupies Mr. Johnstone, with the assistance of IT. Mercer
and a joiner, for about a fortnight, and it occasionally
happens that there are specimens which become accident-
ally broken or damaged, or for some other reason require
to be replaced from the Laboratory, and re-labelling is
sometimes necessary. All this, and the correspondence
connected therewith, has taken up a not inconsiderable
part of our Liverpool Assistants’ time during the past
year ; but I think we are all agreed that it is well worth
doing. The collection, it will be remembered, was
originally formed for the Fisheries Exhibition at the
Imperial Institute in 1897, and as it mainly illustrates

76 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

our local fisheries and local investigations it 1s important
that it should be shown locally, and to the people who are
interested in these fisheries and who contribute to the
expense of regulating them. ‘The educational value of
the exhibition must also be considerable, and may perhaps
be gauged by the keen interest shown by visitors.

I can speak to that from my own experience when the
collections were at Liverpool; and I may also quote a
few sentences from letters sent to me by Mr. B. H.
Mullen, M.A., the Curator of the Museum at Salford
when the Exhibition was in that town, as follows :—

“The Fisheries Exhibition is a very great attraction
here, and must be doing a lot of good. During the past
three days we had almost 1,000 more visitors than during
the same period in 1897. I place the greater part of this
to your most interesting exhibition.”

“You will notice that in those four risen (June,
July, August, peptcurper) last year over 95,000 persons
visited our museum.’

‘“‘T estimate the number of persons who visited the
Museum while your exhibit was here to be 119,852—-say,
120,000.”’

Mr. Mullen prepared, partly from the catalogue given
as an appendix in our last Report, a ‘‘ Popular Guide,”
which was largely sold to visitors to the Salford Museum
at the price of one halfpenny. All this can scarcely fail to
do much good in interesting the public in the importance
of Fishery questions, and in disseminating correct and
useful information as to the work of the Lancashire Sea-
Fisheries Committee.

It is proposed that the Exhibition should be sent in April,
1899, to Warrington, and after that to Bolton, and then
St. Helens, and any others of our neighbouring towns that
can provide suitable accommodation, in the order of their

SHA-FISHERIES LABORATORY. rink

formal application.* The permanent home of the Exhi-
bition, when not on loan, is in the Fisheries Museum at
University College, Liverpool.

It may be mentioned, in this connection, that he May
Professor Boyce and I exhibited at the Royal Society the
practical results of our investigations on Oysters and
Disease which are discussed further on in this Report.

The educational aspect of such matters naturally leads
me to a question of Technical Instruction, which has
arisen almost simultaneously on the Scientific Sub-
Committee and at University College, Liverpool, viz. :—
The formation of a ‘“‘ School of Fisheries Science,” or a
curriculum of instruction in the sciences which underlie
Fisheries knowledge and investigations. In November
the Scientific Sub-Committee drew up the following
report :—‘‘ Having regard to the increasing development
of knowledge on Fishery questions, the Sub-Committee
has considered a scheme for the establishment of a School
of Fishery Science, prepared by Professor Herdman, and
is now in communication with the Technical Instruction
Committee of the County Council, with the view to this,
or some suitable scheme for enabling students to obtain
advanced instruction in Fishery Science, being adopted.
Tt has also been considered desirable, if arrangements can
be made, to allow the Sea-Fishery Bailiffs to avail them-
selves of opportunities for instruction in more advanced
knowledge than they possess of sea-fish, their habits and
food, in order that they may be able to instruct the fisher-
men in matters of which, at present, there is probably
considerable ignorance, and upon which accurate know-
ledge would be advantageous.” This report was approved
by the General Committee at the meeting in November.

* Applications should be sent to Prof. Herdman, University College,

Liver pool.

78 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The matter has also been fully considered by a com-
mittee at University College, which has drawn up a
detailed curriculum of instruction, and the scheme now
only awaits a conference between the University and the
County authorities before becoming a working department
in a most important branch of technical knowledge.
With our extensive scientific Laboratories and the
Fisheries department and Museum at University College,
with our sea-fish hatchery and experimental tanks at Piel,
and with the steamer and system of bailiffs under Mr.
Dawson, we possess already organised and co-ordinated in
Lancashire such a mechanism for instruction in Fishery
knowledge from the scientific, from the industrial, and
from the administrative sides, as probably does not yet
exist elsewhere in the country.

If the County Technical Instruction Committee will
help in meeting the necessary expenses of selected
students passing through a two or three years’ curriculum,
and the Laboratory expenses of showing to the bailiffs and
a limited number of fishermen the methods of investigation
and the microscopic, chemical, and other facts which they
hear about in lectures and reports, it can scarcely be
doubted that much more will be done thereby in the
dissemination of real and accurate knowledge than can
possibly be effected by imperfect and sporadic courses of
lectures to the fishermen.

Turning now to such special work as I have been able
to do myself for this Report :—

(1.) I have considered it useful in the present state of
affairs to discuss somewhat fully Mr. Fryer’s criticism
of Sea-Fish Hatching in America, which appeared in the
last Report of the Inspectors for England and Wales
(1898). Ihave had: some correspondence on the matter
with the United States Fish Commission, and I may

SEA-FISHERIES LABORATORY. | 79

quote here the following sentences from a letter recently
received from the Commissioner :—‘“‘ For about ten years
the Cod work has been attended with marked success,
and in Massachusetts has resulted, not only in establish-
ing the in-shore Cod fishery on grounds long exhausted,
but, through favourable distribution of the fry, in extend-
ing the fishery to other waters not originally frequented
by the Cod.” . . . “Some investigations made a few
years ago by the Commission indicated that the value of
the Cod now annually taken on new grounds is at least
several times greater than the entire yearly expenditures
of the Commission for fish-cultural work, and is increasing
each season.”’

(2.) The investigation into the condition of Oysters from
various localities and under various circumstances, and
their relation to infective diseases in man, which I have
been carrying on during the last three years in conjunction
with Professor Boyce and Dr. Kohn is now concluded,
and our full memoir on the subject with illustrations of
the detailed evidence will soon be published. I have
therefore considered it right to lay before you in a special
section of this report the final conclusions at which we
have arrived on ‘‘The Oyster Question;’’ and I have
appended to it a reprint (from our paper at the Bristol
meeting of the British Association in September) of Dr.
Kohn’s account of the presence of iron and copper in
certain Oysters. I think it is clear that what the public
wants at present is an assurance that the Oysters they
buy and eat come from grounds that are above suspicion.
There is a great opportunity for an independent autho-
rity—either ‘‘ Health” or ‘“‘ Fisheries,” ‘‘ Central” or
“‘ Local **—to take up the matter, and after due investi-
gation to license or certify certain grounds or certain
Oysters upon the lines I have indicated in the conclusions

on p. 184. W. A. HERDMAN,

JANUARY, 1899.

80 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Fish HatcHING WorK AT PIEL.
(ANDREW SCOTT.)

At the beginning of December, 1897, I entered upon my
duties at Piel, and at once proceeded to place everything
in working order for the approaching spawning season.
Various little defects in the machinery and tanks were
discovered and put right. By far the most serious defect —

was found in the large overhead store cisterns in the tank —

room, where considerable leakage was taking place, making
the room uncomfortable to work in. However, with some
difficulty and expenditure of time, they were made fairly
water-tight, and at the end of twelve months are now
practically free from leaks. |

The wooden hatching tanks used in the previous season
were cleaned out, fresh sand put in the filtering compart-
ments, and a constant circulation of sea-water established.
All the apparatus was in satisfactory working order by the
end of January.

On January 27th the steamer visited the spawning
grounds in the central part of the Irish Sea, between the
Isle of Man and Lancashire, where a few hauls with the fish
trawl and surface nets were taken. No mature fish were
obtained, but in the tow-net collections made at the “ Top
end of the Hole”’ and ‘“‘On the Shoals” a few fish eggs
were observed. The grounds were again visited during
February, and although no spawning fish were found, a
marked increase in the number of floating eggs was
noticed. A quantity of living tow-net material brought
into the Laboratory was found to contain three forms
of fish eggs, in various stages of development—in some
the larvee were quite lively. These eggs were carefully
picked out and placed in glass aquaria, with a constant
circulation of sea-water. In the course of the same even-

SHA-FISHERIES LABORATORY. . 81

ing a few of the larvee hatched out, and two forms were
identified, from the arrangement of the colouring matter,
as being the larvee of Cod and Flounder. The third, a
colourless larva, was not identified, and only lived a few
days. The Cod and Flounder larve lived for twelve days.
By this time the yolk sac had been completely absorbed,
but the larvee made no attempt to feed, although kept
supplied with plankton taken in the tow-net.

On March 9th the steamer again visited the spawning
grounds, and this time secured a number of mature Cod,
Haddock, and Plaice, from which a quantity of eggs were
obtained and fertilized. These, however, were probably
not quite mature, and at the end of the following day
they had all died and sunk to the bottom of the tanks,
no development having taken place. The tow-nettings
contained an increased number of developing eggs of Cod,
Flounder, Plaice, &c.

Amongst some living fish brought in by the steamer
from this expedition was a mature female Flounder.
These fish were placed in the tanks, and on the following
day the Flounder was observed to be shedding its eggs.
The fish was therefore taken out, and the eggs pressed
out into a jar, and successfully fertilized with the milt of
a male Flounder that had been captured, amongst other
fishes, in the Barrow Channel by Mr. Wright, a few days
previously. Development proceeded rapidly, and eight
days later all the larve hatched out, there having been
practically no mortality amongst the eggs. The larve
lived without loss for fourteen days. The contents of the
yolk sac had been absorbed several days previously. At
this stage a marked mortality set in, and during the next
few days the larve died off rapidly. Notwithstanding the
various experiments tried to persuade them to feed, at the
end of eighteen days all had died.

82 ‘TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

On March 16th the steamer arrived with a further —
supply of eggs from Plaice and Flounder. None of the —
Flounder eggs, and very few of the Plaice eggs, had been

fertilized. On the 18th March a supply of Cod eggs was
brought in by the steamer. Many of these were fertilized,
and the development proceeded rapidly.

The spawning grounds were again visited on March
22nd, and a number of sailing trawlers were boarded

when hauling in their nets. Many mature fish had been

captured, and a supply of Cod, Haddock, Plaice, Flounder,
and Dab eggs were obtained. The eggs of the Haddock,

Plaice, and Dab did not fertilize, nor did many of the

Cod and Flounder.

The surviving eggs of the Plaice fertilized on the 16th,

of the Cod on the 18th, and of the Flounder and Cod on
the 22nd, and 23rd March developed satisfactorily, hatch-
ing out at the end of eighteen, sixteen, eleven, and thirteen
days respectively. Throughout these periods a consider-
able daily mortality was observed, which was, no doubt,
due largely to the condition of the eggs when fertilized.
In several cases the larve were clearly visible through the
membrane of the eggs when death occurred.

Through the kind permission of the Fishery Board for |

Scotland, the steamer was allowed to visit the Clyde and
trawl there for spawning fish. Three visits were made,
but on only one of these, the first, were mature fish
obtained, and the eggs successfully fertilized. The eggs
were from Plaice, and the quantity on arrival at Piel
measured 150 cubic centimeters. This was by far the

greatest individual quantity of eggs received during the ©

season. On the first visit the eggs of the Witch, Dab,
Grey Gurnard, and Haddock were also obtained; but the
_ Gurnard and Haddock eggs did not fertilize. The Dab

SHA-FISHERIES LABORATORY. : “88

and Witch eges were fertilized, and underwent partial
development, but died after three days.

Development in the Plaice eggs proceeded fhroueh its
usual course with scarcely any mortality, and at the end of
twelve days, the hatching of the larve commenced, and was
completed on the following day. Shortly after hatching
was completed the larve were carefully transferred to
glass aquaria, through which a constant circulation of
water was maintained, and when they were a week old
various experiments were tried to persuade the larve to
feed. To one jar material collected in the filter was
added; to a second, Diatoms; to a third, Copepoda; toa
fourth, plankton from the tow-nets, and to a fifth, ‘‘ Mussel
broth ”’ obtained by squeezing up the living shell-fish and
passing the semi-fluid mass through a fine sieve. All
these experiments were of no avail, and the larve began
to die off when they reached the age of fourteen days.
During the following fourteen days there was a consider-
able daily mortality, and at the end of twenty-nine days
all the larvee were dead, they, apparently, having made no
attempt to feed, as no trace of food could be seen in the
stomach when examined under the microscope.

Other expeditions were made to our own spawning
grounds in search of mature fish, but no more fish eggs
could be obtained.

The method of collecting fertilized eggs by the steamer
is one of some difficulty and uncertainty, owing to the
stormy weather sometimes experienced during the spring
of the year. It may happen that just at the very time
spawning fish are on the ground it is unsafe for the
steamer to venture out. ‘The collecting of eggs from fish
caught in the net of an ordinary trawler is also unsatis-
factory, owing to the circumstance that when the eggs are
ready for shedding the least pressure on the sides of the ~

84 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

fish forces the eggs out. Fish caught in a trawl-net are
generally mixed up with a large quantity of debris,
especially after a four or five hours’ drag, and the weight
of this ‘‘rubbish”’ pressing against the ripe fish forces the
eggs out before the net is emptied on deck. The majority
of the eggs that are brought in to our hatchery from
these expeditions, therefore, are not perfectly mature,
consequently the eggs may not be fertilized at all, or if
fertilized, die off before hatching out. It is from this —
cause that the high mortality arises.

An instance of this was demonstrated during the past
season. A large female Plaice, fully distended with eggs,
was brought in by the steamer from one of the expeditions,
and was kept alive in one of the tanks. In the course of
a few days however, it turned sickly, and to all appear-
ances was in a dying condition. The eggs were therefore
pressed out into a bucket containing a small quantity of
sea-water, and mixed with the milt of a male Plaice that
had been brought in along with the female. None of the
egos floated, although examination showed a number of
them to have been fertilized, and to be undergoing develop-
ment. The living ones were carefully picked out and
placed in a jar by themselves. Notwithstanding a con-
siderable daily mortality, development proceeded rapidly,
and in a few days the little fishes were showing very
clearly through the egg membranes, but only two hatched
out. A number of the embryos reached the hatching-
out point and then died. The two that did hatch out were
feeble in their movements, and lay on their backs at the
bottom of the jar. They were very different from the
larvee of the Clyde Plaice, and only lived a few hours.

It is clearly evident then, that fish eggs for hatcheries
must be under the most natural condition obtainable.
The only way to secure this is to collect the fish towards

SEA-FISHERIES LABORATORY. 85

the end of the previous year, or very early in the spawning
season, and keep them alive in a suitable pond, allowing
them to shed the eggs by their own efforts.

That the water is satisfactory for the purpose is proved by
the result of the past year’s hatching work. After passing
through the filter it was perfectly pure and transparent,
the specific gravity varying from 1:0025 to 1:0026, and the
temperature from 4° C. to 4°8° C.

The mortality of the larve kept after hatching is very
serious. Further experiments will be tried, as it may be
that we have merely not yet hit upon the proper food.
It may, however, be found best to set the young fish free
before they reach the critical stage in their development.

OBSERVATIONS ON THE OCCURRENCE AND HABITS OF
LEPTOCEPHALUS.

(ANDREW SCOTT.)

In recent years much light has been thrown upon this
peculiar group of fishes (the Leptocephalide), which were
at one time considered to be fully developed animals and
classed by naturalists as such, under the generic name of
Leptocephalus. Thanks to the observations made at
Roscoff, in France, in 1886, the form then known only as
Leptocephalus morris, was actually observed to change
into the Conger eel (Conger vulgaris), and later, in 1891-95,
two Italian investigators, Grassi and Calandruccio, carried
on careful experiments on the Leptocephalus brevirostris
taken in the Straits of Messina. They found that this
so-called species went through a transformation, changing
into the common Eel, Anguilla vulgaris.* So that there
can no longer be any doubt now that the Leptocephali of
the older naturalists are only the larval stages of Hels.

* Proceedings Royal Society, London, vol. LX., No. 863. Dec., 1896.

86 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

That there is still a considerable amount of definite
information wanted regarding the movements and true
habitat of these larve in our own seas, may be gathered
from the fact that the records of their capture along the
British coasts are few and far between.

In our surface tow-net gatherings, taken along the
Lancashire coast during the past year or two, we have
occasionally found a little flat, transparent fish, which has
been entered on our lists as ‘‘ Leptocephalus sp.” In
January, 1896, three were taken in the estuary of the
Wyre, by tow-net worked from the steamer. In April of
the same year, one was taken by tow-net worked off
Lytham Pier. These were forwarded to us by Mr.
Ascroft. In January, 1897, one was taken in the Mersey,
off New Brighton, and in the year finished (1898), a
number of individuals have passed through my hands, as
follows :—

On April 26th, Mr. J. Wright, Chief Fishery Officer at
Piel brought into the Laboratory a tow-net collection he
had made in the vicinity of the north end of Roa Island,
and amongst the material was a living Leptocephalus. .
This specimen was kept alive for a few days, but eventually
died. Shortly afterwards, May 18th, while I was collect-
ing young flat-fishes in the shore pools and gutters at low
water in the same neighbourhood, three Leptocephali were
captured. From that date onwards, to the end of June,
when the weather and tide permitted, careful examination
of the shores was made, with the result that eighteen
specimens of this hitherto rare fish were obtained; others
were seen but not captured.

The method adopted for the capture of these and other
young fish, and which proved very successful, was the
following one :—Advantage was taken of the fact that,
during the ebb of the tide here, there is a rapid fall of

SHA-FISHERIES LABORATORY. 87

water, which has scooped out regular well-defined gullies
in the more level stretches of the shore, and although the
adjoining parts of the shore may be almost dry, there is
still a considerable current sweeping down these gullies.
By closing up the smaller gullies, and digging gutters
between them and the larger ones, we were able to divert
almost the whole of the retiring water on certain parts
of the shore into one or two main gullies, which we after-
wards partially closed up with stones, leaving only sufficient
space in the centre to hold a tow-net. On going to these
places during particular states of the ebb, which later on
we were able to locate with considerable accuracy, and
fixing our nets so that the water passed through them, we
were able to secure practically all the material swept off
a considerable area of the shore by the force of the
receding tide. On favourable days, when there was no
wind and the water free from suspended mud, we could
actually see the various kinds of young fishes, Crustacea,
&c., being carried into the net with the current. The
nets were lifted up from time to time, and their contents
emptied into collecting bottles. After the water had
ceased running, we removed the nets and returned to the
Laboratory with our captures, where they were sorted
out, the fish being placed in tanks and glass aquaria.
The Crustacea captured were chiefly used for feeding fish
already in the tanks.

It was amongst the fish taken in this way that we
obtained the majority of our Leptocephalt. Occasionally
others were captured by forcing the tow-net over the
surface mud at the roots of Zostera, which is fairly
common in some parts of our district. With everything
in our favour, we could almost depend with certainty on
having at least one Leptocephalus each time we tried for
them, Sometimes we would get two, and once we cap-

88 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

tured four. At the end of June, when we ceased finding
them, we had as many as fourteen Leptocephali living in
the glass aquaria. They were then measured one by one
and placed in two aquaria, which had a layer of sand on
the bottom. The sizes ranged from 2;% to 2,% inches in
extreme length, =, of an inch in vertical depilh from fin to
fin, and about 74 of an inch thick. They were flat, colour-
less, and perfectly transparent, the viscera, and the heart
and its movements, being easily seen through the skin. -

On being placed in the aquaria, the Leptocephalh swam
swiftly round the sides, with an undulating movement,
like that of the sand-eel. They soon settled down to
their new surroundings, and quickly buried themselves in
the sand. The movements gone through in burrowing
are exactly similar to those made by the sand-eel. The
head is first directed into the sand, then by a rapid back-
ward and forward movement of the posterior part of the
body, the anterior part is forced into the sand, and finally,
by a gliding motion, the posterior part disappears. During
the day-time the Leptocephalt remained hidden away in
the sand. On the slightest disturbance of the water, such .
as would be caused by the throwing in of food, their heads
would be thrust out and a rapid survey taken to ascertain
the nature and position of this disturbing element. If it
were food that happened to fall close to them, they would
seize it without coming entirely out of the sand, and
would then glide backwards into the burrow. If the food
did not fall within reach, they would not venture to pur-
sue it in daylight. On going into the tank-room at night,
when all was in darkness, and suddenly flashing a light on
the aquaria, it would usually be seen that the Leptocephalt
were swimming about actively, but soon retired into the
sand if the light were continued.

The Leptocephalt were kept under observation for a

SHA-FISHERIES LABORATORY. 89

week or two, when all, with the exception of two, were
unfortunately lost through the overflowing of the aquaria
in the night, no doubt having jumped out when the jars
were full. The survivors remain alive, and at the end of
November measured 375 inches in length, one having
grown three-fourths of an inch in four months and the
other slightly less. The transformation of these larve
from the Leptocephalus stage was not actually observed,
but on June 30th they were flat, transparent, colourless
Leptocephali; and on August 3rd had passed into young
eels, having a smoky-tinted back, silvery-grey sides, and
being no longer transparent.

During the earlier parts of August numbers of young
eels were found under the stones on various parts of the
shore, which exactly corresponded with the appearance of
the transformed Leptocephal.

From these observations it would thus appear that
the Leptocephal are inhabitants of the mud, and their
occasional presence in surface tow-net collections is due
to their having been swept out of their burrows by the
strong currents, and that they are never taken in the
dredge is owing to their activity. Attempts to capture
them with a tow-net when they are swimming against the
current usually ends in failure.

In passing, it may be noted that all the Leptocephali I
have obtained from tow-nettings on the Lancashire coast
are identical with the hemi-larval stage described by
Grassi, and many of the observations agree with those
made by him, and published in his paper in the Proceed-
ings of the Royal Society already referred to.

‘90 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

OBSERVATIONS ON THE HapBits AND Foop oFr YOUNG

FISHES.
(ANDREW Scort.)

Early in May the young flat fishes hatched out under
natural conditions in the sea began to appear on all the
sandy shores. By allowing the water draining down the
sullies, when the tide was ebbing, to pass through a tow-
net, considerable numbers were captured. Many of these
were kept alive in our tanks and aquaria; some were pre- -
served for future study, and others were examined at once
to ascertain the nature of the stomach contents.

These little fish, chiefly Plaice and Flounders ranging
from two-fifths to three-fifths of an inch in length, were
quite colourless and transparent, the stomach and alimen-
tary canal showing clearly through the skin. Although
they had assumed the flattened character of their parents
and the eye had begun to move over the head, they still
swam about in a vertical manner. In the course of a day
or two they were observed to have considerable difficulty
in maintaining the upright position, and ere long, after a
few more feeble attempts to swim vertically, they settled
down to a semi-sedentary life in the sand. During the
day-time they remained buried, except the mouth and eyes,
and could only be detected with difficulty. In the dark-
ness they came out and swam freely on the surface of the
sand. After a few weeks, when the little flat fishes had
become accustomed to their surroundings, they ceased
burying themselves, and simply lay on the surface. Some-
times they clung to the sides of the jars with great
tenacity.

The food of these young fishes was found to consist
almost entirely of Copepoda. Collections made later on,

SHA-FISHERIES LABORATORY. 91

when the fish had grown to an inch and upwards in
length, showed them to be feeding on Mysis alone.

From their peculiar structure, one would naturally
expect that the flat fishes would be rather sluggish in
their movements, and not at all particular as to the nature
of their food. Far from this being the case, however,
they pursue their food with much vigour and select a
special diet, as 1s clearly shown when one examines the
contents of their stomachs. The stomachs of the smaller
ones, from one inch up to four inches in length, captured
on the shores of our neighbourhood, are usually almost
entirely filled with Mysis, a group of Crustacea that depend
chiefly on the power to escape capture by making sudden
leaps when approached by any moving object. The flat
fish appear to be aware of this peculiarity, as they care-
fully stalk the Mysis, and when they get close up make a
sudden spring, seldom failing to capture their prey. That
the young flat fish prefer living to dead food can easily be
seen by throwing a mixture of dead and living Mysvs
amongst them. ‘The fish are always on the look out for
- food, and at once proceed to investigate any object that
makes its appearance in their vicinity. If the Mysis
swims or leaps away, then it is pursued and captured, but
if it makes no attempt to escape, the fish will abandon it
for a more lively prey. Of course, when the fish are
hard pressed for food they may not be so particular in
waiting until the object shows considerable signs of life
before they capture it.

The older flat fishes, from four inches and upwards,
captured on the Roosebeck Scars, usually feed on young
shelifish, such as Mussels, &c., worms (Arenicola), and
Crabs (Carcinus).

Shortly after the advent of the flat fishes, the young of
the various round fishes, such as the codling, bluffin, sand-

92 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

eel, herring, goby, lumpsucker, stickleback, and lesser
grey mullet, began to make their appearance in regular
order; each having their own manner of capturing their
prey, and all, with the exception of the young mullet,
feeding on Mysis. Even the ungainly-looking and
awkwardly-swimming young lumpsuckers are able to
capture Mysis. They swim after their prey at, for them,
a rapid rate, making sudden dashes as they pass, and
usually trying to seize the Mysis by the middle. This
is probably done to prevent too much attention being
bestowed upon them by their less fortunate companions,
which they would otherwise be sure to receive if the prey
were captured by the head or tail. It occasionally happens
that the young lumpsuckers do capture the Mysis by one
end, and before they can swallow it another lumpsucker,
usually a smaller one, has seized the free extremity. It
is rather an interesting sight to see the stronger one try-
ing to shake off the weaker, but so tenaciously do they
cling to their victim, that the smaller fish is frequently in
danger of disappearing after the Mysis. Only at the last
moment does it reluctantly relinquish its hold.

The last of the young fishes to appear on the shores in
the vicinity of Piel, so far as has yet been observed, are
the lesser grey mullet. Numbers of these fish were
captured in a well-defined gully on the east side of the
breakwater joining Foulney Island to the land. ‘They
were first noticed about the middle of September, and
were then fully an inch in length. The stomachs, on
examination, were found to be filled with vegetable food,
chiefly Diatoms, Navicula being the prevailing species.
Later on, when they had reached the length of one and a
half inches, the food was found to consist of a mixture of
Diatoms and Copepoda (Tachidws).

There appears to be little or no difference in the food

SEA-FISHERIES LABORATORY. 93

of the young fishes frequenting the shore, and no matter
whether they were captured at mid-day or mid-night, the
the food was always the same. On several occasions we
trawled the gullies at mid-day and mid-night with a small
otter shrimp net. This was found to be very successful
in capturing young fish, &c.

A number of experiments were made to ascertain how
far the colours of certain Crustacea protected them from
falling a prey to the fishes. It was found that when semi-
transparent and dark-coloured Mysis were put in the jars,
the colourless ones were eaten before the dark. Similarly,
when a large number of variously coloured Hippolyte (Vir-
bius) varians, ranging from transparent to almost. black
were used, the transparent ones were the first to disappear.
Gradually the others were captured; last of all, but very
seldom, were the dark ones pursued. Are the pigmented
forms less noticeable under the circumstances, or is Ene
pigment itself distasteful ?

THE PLANKTON WORK.
(ANDREW SCOTT.)

The examination of the floating plankton collected in
the vicinity of the Lancashire coast has been continued
throughout the greater part of the year. A satisfactory
investigation of the material, and the accurate identifi-
cation of the organisms contained therein, is a matter of
considerable difficulty, owing to the large quantity of
debris that is nearly always present.

Areas of the sea into which the contents of large rivers
flow are usually contaminated with the spoil carried off
the land. This finds its way into the river either by
accident or intention, and the period during which the

94 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

lighter particles remain afloat depends largely upon the
strength of the current that carries the material along.

Practically the whole of the Lancashire coast-line is
influenced more or less by the outpouring of large rivers,
such as the Dee, Mersey, and Ribble in the southern and
central part, and by all the rivers flowing into Morecambe
Bay in the north. Therefore, unless we can go to a con-
siderable distance from land, our tow-net collections
are almost wholly composed of vegetable debris, the
land origin of which is clearly demonstrated by the
presence of sporangia of ferns, seed capsules and leaves
of mosses, protecting scales of leaf-buds, twigs and leaves
of trees, etc.

The quantity of debris present in these local tow-
nettings is, of course, subject to weather and tidal
influences. After a spell of calm, during neap tides, we
occasionally get a gathering nearly free from rubbish.
One or two gatherings from the vicinity of the Bar Light-
ship in the Mersey consisted almost entirely of the
Copepod Hurytemora. In the summer of 1895, gatherings
were taken in the Rock Channel, which contained nothing
but large quantities of Nocteluca. This was unusually
abundant throughout the southern part of our district for
a few weeks, occasionally giving the water a distinct
brown appearance. None of the gatherings taken in the
Barrow Channel, outside of Walney Island Lighthouse,
have been free from debris, and the same is true of those
taken in the Ribble, off Lytham Pier, and in the neigh-
bourhood of Nelson Buoy. Mr. Ascroft’s system of
placing his tow-nettings in white dishes with clean sea
water, and allowing the organisms to separate out and
come to the sides of the vessels, where they are secured
and preserved, is a very useful one, but, unfortunately,
not always practicable by our fishery officers, as their
police work takes up most of their time.

SHA-FISHERIES LABORATORY. r 95

After the experience gained last year, we arranged that
the gatherings should be taken in more seaward positions
during 1898. This has been done, but the results are
very much the same as before. The weather is frequently
quite unsuitable for our sailing boats to venture far from
land, and on calm days rowing out to the stations is
tedious and dangerous, owing to the tides.

To gain an accurate idea of the floating plankton of the
Irish Sea other means will require to be adopted. The
method that suggests itself as being the most convenient
from all points of view, is to make use of the lightships
that are anchored off various parts of our coast. There
is probably sufficient current set up by the rise and fall of
the tide to keep a tow-net extended. By supplying the
keepers with bottles containing preservatives, tow-nets,
and necessary instructions for working the nets, pre-
serving the material collected, and the times (night or
day, or both) when the collections should be taken, a
more satisfactory knowledge of the plankton would be
obtained. One of the men when off duty might be taken
out in our steamer and shown the methods. Then the
steamer could visit the lightships when convenient, say
once a month, leave a fresh supply of bottles, replace
worn out tow-nets, and bring back the collections taken
during the interval.

The success of this method, of course, depends entirely
upon (after the necessary permission has been obtained)
the zeal and care of the men themselves, and the en-
couragement we give them.

The collecting stations which might be tried this
ensuing year are the North-West, the Morecambe Bay,
the Selker, and the Bahama Lightships, and if the results
prove satisfactory, which no doubt they will, the system
can easily be extended. :

96 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The material examined during the past year has given |
us results which vary very little from those already given
in former Reports. The following are a few of the more
noteworthy organisms observed :—

Mysis occurred in many of the gatherings throughout
the year, but only a few individuals were present in each
tow-netting. They were very common during the
whole year in the shore pools in our neighbourhood. On
September 14th, immense number of young Mysis were
observed in the channels near Baicliff. This is un-
doubtedly one of the most valuable sources of food supply
for young fishes.

Crangon, Pseudocuma, Gammarus, and Hurydice were
occasionally noticed, but only one or two individuals at a
time. Ona warm day, when the sea is calm, numbers of
Hurydice may be seen disporting themselves on the
surface. In their movements they are not unlike the
‘‘ Whirligig” beetle of the fresh-water ponds.

Copepoda—EHurytemora, Acartia, Paracalanus, Temora,
Calanus, Pseudocalanus, and Ovthona, were present more
or less throughout the year, but never, as a rule, in any
quantity. In a few of the gatherings from the vicinity of
the Bar Lightship, Hurytemora was, on one or two
occasions, very common. The striking difference between
local and off-shore collections was clearly demonstrated
by the comparative scarcity of Copepoda in-shore. As
early as January 10th one-fifth part of the tow-nettings,
taken by the ‘‘ John Fell,” five miles north of the Selker
Lightship, consisted of Calanus, Paracalanus, and Acar-
tia. At the same time, only a few individuals were taken
in local gatherings. ;

At a very early stage in the life history of most fishes,
Copepoda play an important part as a food supply. The
majority of the young flat fishes—Plaice, Flounder, Dab,

SRA-FISHERIES LABORATORY, ae

&c.—when they first appear on the sandy shores, feed
almost entirely on these minute crustaceans. The species
usually found in the stomachs of the young fishes, between
half-an-inch and an inch, are chiefly littoral, such as
Eurytemora, Ectinosoma, Tachidius, and Jonesiella. Al-
though the Copepods at particular stages form a food
supply for the fishes, it is just possible that the fishes
themselves, when they are newly hatched, may be eaten
by the Copepods. Instances of this were demonstrated
during our feeding experiments, and a species of Copepod
(Centropages) was seen to capture one week old Flounders
and eat them.

Other invertebrates, such as Sagitta, Meduse, Cteno-
phora, and Ozkoplewra occurred very sparingly in the
local gatherings. :

The eggs and larve of fish were very plentiful in tow-
nettings taken in the open sea, but very few eggs and no
larvee occurred in the local collections. The first collec-
tion in which fish eggs were observed were those taken
by the steamer on January 27th, at the “Top end of
the Hole.’”’ None were found in local gatherings until
February 4th.

Amongst the surface material collected by tow-net when
the steamer was in the Clyde on April 2lst, were three
large fish eggs, measuring fully three millimeters in
diameter. ‘The larvae were well developed, and hatched
out a few days later, but did not survive. There was one
large and one small amber-coloured oil globule present,
but no space between the embryo and the egg membrane.
The species of fish to which these eggs belonged is still
uncertain. The only eggs corresponding to them in size
are those of the Halibut; but the eggs of this fish have
apparently not yet been taken in surface nets. The
newly hatched larve had a large yolk sac and very short tail,

98 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Larve of various groups of the invertebrata, including
Mollusca, Crustacea, Polyzoa, Vermes, and Ccelenterata,
were occasionally observed in the local collections. The
floating eggs of Alcyoniwm were very common in Barrow
Channel, off Roa Island, in the beginning of April, and
also in the tow-nettings taken in the Clyde.

Noctiluca was very common in the collections taken in
the vicinity of the Bar Lightship, Mersey, on June 24th
and onwards; but was not observed in the northern part
of the district until the end of August.

Ceratium tripos, C. furca, and C. fusus, occurred in
many collections throughout the year.

Diatoms (Biddulphia, Coscinodiscus, &c.) were very
seldom found in the local collections, but were abundant
in the gatherings made by the steamer further out, in the
spring months.

EXPERIMENTS WITH WEIGHTED Drirt BOTTLES.
(ANDREW SCOTT.)

In a former Report,* under the title, “‘ The Drift
Bottles and Surface Currents,” in which the result of the
distribution of drift bottles over the Irish Sea, and the
conclusion drawn therefrom, is given, reference is made
to an experiment of Mr. Ascroft’s, where, instead of using
small bottles containing only the post card, larger bottles
were employed, but, in addition to the post card, were
so weighted with sand that they floated almost entirely
submerged, and of which nearly 30% were returned. It
is also incidentally remarked that some of them sunk out
of sight when set free, one being subsequently brought up
in a steamer’s trawl while fishing in the vicinity of the
Bahama Lightship on the north-east coast of the Isle of

* Report for 1895, Sect. II., p. 12,

SHA-FISHERIES LABORATORY. : 99

Man. This is now accounted for by the following explana-
tion. During transit some of the bottles were broken,
but in order that the post cards should not be wasted,
other whole bottles were obtained, the card and the same
sand put in, then the whole sealed up, over-looking the
fact that all bottles may not be of the same weight, each
requiring careful ballasting, consequently when these
bottles were thrown overboard the majority of them
sank.

The result of this experiment of Mr. Ascroft’s showed
that weighted bottles tended to go south, thus differing
from the light ones, which largely went to the north. In
order to throw further light on this apparent southerly
drift of weighted bottles, it was decided to give the
weighted bottles a further trial, and in addition, by set-
ting them free on the spawning grounds when the steamer
was trawling for spawning fish, perhaps gain some definite
information regarding the direction of the drift of the
‘surface waters in which the embryos and larval fish
usually are.

Accordingly we had a number of ordinary post cards
having the same notice previously used, printed on the
back, and we purchased a supply of bottles, known as ‘‘ cocoa
wine bottles,” of about one pint capacity. These bottles
were placed one by one floating in a bucket of sea-water
of 1:0026 specific gravity, and then carefully ballasted with
dry sand, so that when the rolled up and numbered post
card was placed inside, the cork inserted, and the whole
sealed up with paraffin wax, only about an inch of the
neck of the bottle was above the surface of the water.
They were then sent on board the fisheries steamer in
batches, each batch being accompanied by forms having
the numbers of the bottles, and spaces for the insertion of
the position of the steamer when the bottle was set free,

100 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the date, time, state of the tide, direction of the wind,
and approximate quantity of floating eges taken in the
tow-nets. The forms were filled up by Captain Wignall,
and returned to me, after the bottles had been set free.

Altogether 102 bottles were made up, but one of these
was broken in transit, so that during the time the steamer
was trawling for spawning fish, from February to the
middle of May, 101 bottles were set free, of which we
have exact particulars regarding their distribution. Of
this number 41 or 40°5% have been returned to Piel from
various parts of the coast-line of Cheshire, Lancashire;
Cumberland, and the South of Scotland.

The following table gives the position of the steamer
when the bottles were let off, the place where they were
subsequently picked up, and the number of days that
elapsed between their despatch and recovery (see p. 34) :—

From this table it will be seen that nearly 22% of the
bottles have drifted in an easterly or southerly direction
after being set off, and nearly 18% have taken a northerly
direction. Only one bottle, No. 60, appears to have
crossed the head of the tide. Of the 40 set free in the
vicinity of the Bahama ship, 12 have been returned, and
of these eight have gone north to the south coast of
Scotland, and four have taken a south-easterly direction,
landing on the Cumberland coast. Bottles 51 and 53, set
free within fifty-five minutes of each other, yielded rather
peculiar results, the first going to Holyhead in seven
days, and the second to Duddon in 47 days, distances of
fifty-five miles and twenty miles respectively in a straight
line from the point of despatch, but in exactly oPpaaae
directions.

Of the various days on which the bottles were found,
only three were Sundays, and seven were Mondays. The
conclusion naturally drawn from this evidence is, that the

_.SEA-FISHERIES LABORATORY. 101.

bottles were probably found almost immediately after
being stranded. The shores are so regularly patrolled by
the people living in the immediate neighbourhood, that —
theré is little chance: of any unusual object: being long ©
over-looked. One or two of the finders stated that the ©
bottles came ashore during the previous tide, and one -
was picked up at sea, five miles from Blackpool, by a
fisherman, after it had been 23 days in the water.

In all probability the bottles would float slightly higher —
in the water of the open sea than they did in the bucket —
at Piel, as the specific gravity of the water at the various |
stations ranged from 1°00268 to 1:0027, so that there would .
be little chance of their going towards the bottom of the
sea till they entered the estuaries of rivers.

The table shows that out of the 101 bottles set free, |
fully 30°6% have been stranded on the Cheshire, Lanca- :
shire, and Cumberland coasts, and therefore the result of
this experiment, even with weighted bottles in place of
light ones, confirms, in a striking manner, the conclusion |
arrived at by Professor Herdman* when summing up the .
results of the first series of experiments, 7.e., ‘‘ That the
embryos of fish spawning in the deep water on the eastern _
side of the Isle of Man would go to supply the nurseries _
in the shallow Lancashire and Cheshire Bays.” |

Fish eggs and larvae were found in all the tow- nettings —
made during the experiment. |

[Table over page.

* Rep. Lane. Sea-Fish. Lab., 1895, pp. 20—21.

102 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

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104 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

THE SPAWNING OF THE MussEeL (Mytilus edulis).
| (J. JOHNSTONE.)

During the last year an investigation of the reproductive
organs of the common Mussel has been made in relation
to the period during which spawning takes place on the
beds along the Lancashire sea coast, and to the histo-
logical changes accompanying the ripening and extrusion
of the reproductive products. The methods employed
were :—(1) The microscopic examination of the gonads of
specimens taken by Mr. A. Scott from the Roosebeck scars
and the beds in the Barrow Channel, and of specimens sent
by. the bailiffs from the Wallasey, New Brighton, and
Morecambe beds; (2) the search for free-swimming and
fixed larve on the beds themselves, and (8) the examina-
tion of the in-shore tow-nettings taken by Mr. Wright in
Morecambe Bay, and by Mr. Eccles outside the estuary of
the Mersey. The records of a continuous weekly series of
tow-nettings, taken in the year 1895 by Mr. R. L. Ascroft,
at Lytham Pier, have also afforded valuable evidence.

As a result of this year’s observations, 1t has been found
possible to fix approximately the date of a maximum in
the spawning of the Mussel, during which a rapid and com-
plete extrusion of the genital products, accompanied by
other histological changes in the mantle and visceral mass
of the animal, takes place. In the year 1898 this was
found to begin about the beginning of July and last till
about the beginning of August; but it is probable that the
limits of this period are variable to some extent. There
is, however, considerable doubt as to whether this is the
only time in the year during which spawning takes place,
and various observations render it at least possible that
there is a secondary spawning period early in the year,
aud that there is a continual but slow emission of ova and

SHA-FISHERIES LABORATORY. 105

spermatozoa from the time when these have accumulated
in considerable quantity in the gonads, that is to say, from
the beginning of April on to the beginning of the summer
spawning period. And it seems certain, considering the
variability observed in the ripening of the gonads, that
isolated individuals may undergo complete spawning a
considerable time in advance of, or later than, the date of
occurrence of the maximum. However that may be, there
can be no doubt that the number of larve resulting from
spawning in the early part of the year bears a very small
proportion to those produced during the maximum spawn-
ing period in the summer months.

There is still some uncertainty s caihdition: the rate of
erowth of the young Mussel, and this is due probably to
variations contingent on the conditions under which the
adult animal spawns, and the larva undergoes its early
development. Most probably in the early stages during
which the young Mussel has-a free-swimming existence,
the development and rate of growth are fairly constant,
but with the acquisition of the byssus and the fixation of
the larva, the subsequent growth is dependent, to a large
extent, on the situation it finds itself in, the supply of food,
the extent to which the larve are crowded together, and
on the time of year in which spawning of the parent
occurred. From the observations made by Wilson,* who
succeeded in artificially fertilizing eggs of the Mussel, and
tracing out the early development, it appears that the
larva, about 0°15 mm. in length, provided with semicircular
valves, showing the first rudiment of the anterior adductor
muscle, and still using the velum as a locomotive organ, is
at most 12 days old. An older stage than this, with the

* J, Wilson.—‘‘On the Development of the Common Mussel.” Annual
Report of the Fishery Board for Scotland for the year 1885, pp. 218—222, and
Report for 1886, pp..247—256, Plates XII.—XIV. ;

106 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

valves assuming an oval form, the foot developing, the
posterior adductor muscle present, rudiments of four gill
filaments appearing, and the velum still functional is, at
the outside, a month old. Subsequently to this (0°28 mm.
long) blue pigment is deposited round the margin of the
shell, the foot becomes the organ of locomotion, and the
larva prepares for fixation by the development of byssus
gland and byssus.

Free-swimming larve, somewhat younger than the stage
last referred to, that is, about 0°25 mm. in diameter, and
provided with circular valves showing no trace of pigment,
were taken by Mr. Scott in a tow-net gathering at
Piel, on September 9th, last year. These Mussels were
certainly less than one month old, and their appearance at
this time is in accordance with anatomical observations
made on specimens taken in the neighbourhood, which led
us to fix July and early August as the months during
which spawning took place. But the evidence given by
other observations of this kind is very perplexing. Thus
young Mussels with circular or oval valves ranging in size
from 0°27 to 0°45 mm., and with the rudiments of four or
more gill filaments present, were taken in June, 1892,
fixed to various zoophytes, and assuming the age of these
not to exceed a month, the time of spawning is thrown
back to May or early June. An examination of the
in-shore tow-nettings shewed that young Mussels of
approximately this size (0°3 to 0°6 mm.) were taken in the
estuary of the Ribble on January 27th, 1898, and others,
varying in size from 0:2 to 0°7 mm., were found in Ulver-
ston Channel on February 4th. Some of these had
probably been already fixed and were loosened by the
force of the tide, but others had all the characters of the
free-swimming stages described above, and their presence
at this time is only explicable on the assumption that

S$EA-FISHERIES LABORATORY. 107

there had been considerable retardation in the rate of
growth, or that they had resulted from spawning early in
the year. The early maturity of the spermatozoa is in-
teresting in this connection. In some specimens taken
in December by Mr. Scott, ripe spermatozoa, which
remained alive for about 12 hours, were expressed from
the mantle lobes. An attempt was made to bring about
artificial fertilization, but although the eggs were found to
be covered with motile spermatozoa, and in some cases the
formation of the first polar body took place, the segmen-
tation of the ovum was not observed.

The assumption that some Mussels at least may spawn
early in the year is necessary to explain the presence of
larve in January and February, if it should be found
impossible to account for the presence of such by con-
siderable variation in the early development and rate of
growth; and that there is a gradual emission of spawn
during April, May, and June seems probable also in view
of the fact that free-swimming larve are to be found in
May and June. But from the anatomical standpoint,
such secondary spawning periods are accidental, and do
not effect the statement that there is a yearly cycle of
changes in the reproductive organs of the Mussel, which
begins with the termination of the act of spawning some-
time in the summer months, includes the gradual and
continuous ripening of the gonads, and ends with a com-
paratively rapid extrusion of the reproductive products,
leaving the animal in a ‘‘spent”’ condition, after which
a short period of rest occurs, and the cycle of changes is
repeated. The duration of the maximum spawning period
we have not precisely made out, and its date is probably
variable and dependent on changing meteorological con-
ditions, but that it occurs during May, June, July, and
August seems perfectly certain. ‘This cycle of changes
will now be considered.

108 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

It will be remembered that in the common Mussel. the
sexes are separate. Hermaphrodites, if they exist, must
be extremely uncommon as, in the examination of many
hundreds of specimens, none have been observed. There
is a slight preponderance of males over females, the ratio
in 218 specimens examined being 118: 100; but it is
probable this is too low, as it is hard to avoid selection in
choosing the specimens for examination.* Except fora
slight difference in the colour of the mantle lobes there
are no external sexual characters.

The gonad (Pl. L, fig. 2) is situated in the visceral
mass and mantle lobes; the organ is paired and bilaterally
symmetrical, and consists of a branching tubular gland,
the efferent duct of which is situated on the summit of a
‘small papilla (Pl. I., fig. 1, pp. gen.), about 1 mm. in
height, which lies posterior to the foot and immediately
beneath. the gills (67. &.), and can be easily seen by
reflecting the latter and the mantle (Mn. #.). This duct
is richly ciliated for some distance internally, when the
ciliated wall is partially replaced by germinal epithelium.
Strips of columnar ciliated epithelium (Pl: IL., fig. 7, ep.
cil.) are present along the greater length of the tubules,
but the terminal portions are entirely lined by germinal
epithelium. Passing inwards these tubules branch

* Since the above was written the tabulated results of the examination of
the specimens sent to the Laboratory by the Bailiffs have been referred to.
The portion of these records dealt with relates to the examination of a large
number of Mussels taken at random from most of the Mussel beds in the
district, and sent to the Laboratory in lots of a dozen each, and extends over
the years 1892-96. In order to eliminate as many sources of error as may be
possible, all those sheets in which the sex of one or more of the specimens is
regarded as doubtful, and also those dealing with Mussels taken about the
time of the ‘‘ spent” period, have been rejected. This leaves a total of 821
specimens of which the sex was probably accurately determined. Of these
-449 were male and 372 female, giving a ratio of males to females equal to
6 : 5 (very nearly).

~

SEA-FISHERIES LABORATORY. 109

repeatedly and in quite an irregular manner. Anastomoses
between the various branches have been figured by Wilson,
but we have not seen these. All along the course of the
main branches small twigs are sent out, some no longer than
their own diameter, and these grow and invade whatever
portions of the body they can find room in. The gonad
thus occupies no particular part of the body of the animal
except that the position of the external opening is con-
stant, but it may be ‘conveniently said to be situated
posteriorly to the digestive gland, and as ripening proceeds,
to invade every part of the body, where only connective
or parenchymatous tissue is found (PI. L., fig. 2, twb. ov.).
Simultaneously with the encroachment of the genital
tubules, this tissue undergoes absorption or degeneration.
The keel-like mass, the ‘‘abdomen,” (PI. L., fig. 1., abd.)
situated medially and posteriorly, is thus completely filled
up by the organ, and as maturation goes on, the whole
space in the mantle lobes (Pl. I., fig. 2, Mn. L.; Mn. &.),
between the internal and external epidermal surfaces is
filed up by the branching tubules of the gonad and
except for the merest trace of connective tissue contains
nothing else. Long before spawning occurs these tubules
have invaded almost every part of the body (PI. L., fig. 2),
even the delicate membrane forming the external wall of
the pericardial cavity being overspread by them and
becoming opaque. With the act of spawning the whole
appearance of the animal may be changed, owing to the
emptying of these tubules of their contents, and the
mantle lobes may lose their thick and opaque appearance,
and become thin and transparent. This is the case in
the Mussels in some of the beds in the district, but more
usually the mantle lobes remain thick and opaque. After
spawning it is only by microscopic examination. that the
condition of the animal can be determined. —

110 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

As indicated above, the cycle of changes in the gonads
is a yearly one, or approximately a yearly one, variations
due to the nature of the season doubtless taking place.
This cycle of changes may be represented by four stages.
In giving the dates, we refer more particularly to what was
found during the year 1898. |

STAGE I.—The end of July and the beginning of August.
The Mussel is ‘‘ spent,” that is, the reproductive products
have been entirely extruded, and in the mantle lobes and
visceral mass the genital tubules are, to a large extent,
collapsed or degenerated. The state of the animal in
respect of its reproductive function is one of rest.

STAGE II.—September. The gonads have begun to
invade the mantle lobes and other parts of the body.
Proliferation of ova and spermatozoa from the germinal
epithelium is in progress, and goes on slowly and continu-
ously from now till early in the following year, when it
becomes very active, and the mass of the gonads becomes
much greater.

Stace III.—April. The mantle lobes have attained
their maximum of thickness, and are completely filled up
by the gonads. The tubules composing the latter have
increased greatly in sectional area, and are completely
filled up by ova or spermatozoa in female and male respec-
tively. Proliferation from the germinal epithelium is not
now so active as in the earlier part of the year, and
maturation of the genital products probably goes on.
Where the ova or spermatazoa are in contact with the
ciliated portions of the tubules, they are probably being
swept away and removed out of the body. |

Stace [V.—July and early August. Spawning in the
sense of a complete removal of the genital products is
now in progress. There is a rapid decrease in the mass
of the gonads, both in visceral mass and mantle lobes,

SHA-FISHERIES LABORATORY. Slee

and a corresponding increase in mass of the parenchy-
matous tissue lying between the tubules. Many of the
branches of the tubular gonad disappear entirely, but
many others, and particularly the ciated, non-glandular
portions, persist a sitw. At the end of this period, when
the animal again enters on Stage I., the parts of the body
formerly occupied by the swollen genital tubules are now
the seat either of a massive syncytial tissue, or of a delicate
reticulum, the difference depending probably on the con-
dition of nutrition obtaining on the bed from which the
specimen was taken..

We may now consider the characters presented by the
animal in the various stages referred to more particularly.

STAGE I. (Pl. IL, figs. 3, 4, 6).—Externally there may
be nothing to indicate that the specimen under examina-
tion has spawned, although in a section, or in a cleared
preparation, the difference between this stage and the
preceding one is striking. The sex of the animal is
determinable only with some difficulty, by the presence of
stray masses of ova or spermatozoa, which have failed to be
extruded. Where the germinal epithelium can be recog-
nized, it is not very different in male and female. The
area in a section of the mantle now occupied by the
tubules of the gonads is so greatly reduced that, on a
superficial examination, it might be thought that the
latter had completely disappeared. But more careful
scrutiny reveals the presence of many tubules in a col-
lapsed condition (Pl. II., figs. 3, 4, twb. ov.), the walls
pressed against each other by the pressure of the reticular
tissue. It is, however, only the larger tubules which so
persist ; the finer branches have been absorbed or broken
down in some way.

The space occupied in the ripe Mussel by the gonads is
now filled up by a large celled parenchymatous tissue

112 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

(Pl. II., fig. 8, vet.) of peculiar characters. This tissue
which, in the ripe Mussel, was represented by a few
delicate fibres, lying wedged in between the tubules, has
now increased greatly in mass, and occupies almost the
whole thickness of the mantle lobes. It does not, like the
corresponding tissue in the Oyster, consist of cubical or
polygonal cells, with clear bodies arranged to form a
coarse network, the bars of which are composed of distinct
cells, but is rather a syncytium, in which cell outlines
cannot be readily distinguished. The surfaces of attach-
ment of the parts to each other are large, but occasionally
delicate fibrous connecting strands may be found. The
cells are multinucleated, an isolated patch may be seen to
possess several nuclei, but no trace of cell walls. The cell
substance stains deeply with eosin. In moderately thick
sections it appears to be dotted over with small, light,
circular areas, but in thinner parts these are seen to be
vacuoles, possibly filled with a substance which does not
readily stain.

The epidermis (Pl. IIL., fig. 3, ep. ext.) of the mantle is
formed of a single layer of cells, with large nuclei and
rather indistinct cell walls. On the internal face of the
mantle this epidermis is ciliated. Beneath it there may
be a layer of longitudinal fibrous tissue, but this is not
constantly present. The reticular tissue of the interior
of the mantle is connected with the epidermal layer by
delicate strands, but occasionally large rounded masses
may be seen lying beneath the former. The epidermis
contains eosinophilous cells forming projections internally
and externally. Blood vessels are abundant along the
external face of the mantle (Pl. IL, fig. 4, dl. sp.). The
larger of these have a fine endothelial lining the structure
of which is difficult to make out. These, as well as the
interspaces of the reticulum beneath the epidermis, may
be crowded with blood corpuscles. ;

SEA-FISHERIES LABORATORY. 113

The characteristic of this stage is the enormous develop-
ment of this reticulum, which has all the characters of a
tissue produced by the rapid multiplication of nuclei,
without a corresponding differentiation of cell bodies and
walls. In some sections the space occupied by it is so great,
that the appearance is almost that of a TENE STONE
nucleated matrix, perforated in many places.

The description given of this stage applies to the
Mussels obtained from the Wallasey and Morecambe
beds, the Roosebeck outer scar, and from those in the
Barrow Channel. Mussels obtained from the Roosebeck
inner scar have different characters in respect of the
condition of the gonads at this stage, and these are
probably correlated with the different conditions of nutri-
tion obtaining on this bed, which have been commented
on in a former Report.* Spawning in these Mussels
leaves the mantle lobes delicate and transparent. The
reticular tissue has the form of an attenuated network of
large mesh, and, in consequence of this, the remains of
the genital tubules are more easily seen, and a resting
stage is not so conspicuous.

sTace Il. (Pl. I1., fig. 7; Pl. L, fig. 2).—The period
of rest which follows immediately after spawning, and
which is represented by Stage I., is of short duration, and
as early as the beginning of September the developing
gonads may be seen in the mantle. There can be no
doubt that, after spawning, many of the tubules break
down and disappear, but it seems quite clear that the
more important branches persist in the mantle through
the period represented by Stage I. Towards the end of
September ripening has commenced, and proliferatiom
from the germinal epithelium is in progress. At any

* A. Scott. ‘‘ Mussels and Mussel Beds.” Lance, Sea-Fish. Laby. Report
for 1895, pp. 21—32, hs

114 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

stage this process is further advanced in the male than in
the female. During September the tubules are of small
diameter, no greater than that of an interspace in the
reticulum. The structure of the cells composing the
germinal epithelium of this stage is nearly similar in both
sexes. An irregular layer of cells is supported on a base-
ment membrane. The female germinal epithelium (PI. IL.,
fig. 7, ep. ger.) can be distinguished by the large nuclei of
the cells. At this stage it consists of few cells, and the
first few ova separated off fill the lumen of the tubule.
Proliferation is a more rapid process in the male, and in
a short time the tubules -are filled with cells, resulting
from the divisions in the germinal epithelium. Most of
these cells become spermatoblasts, and divide subsequently
to form the spermatozoa, but many of those separated off
at this stage become modified to form the system of sup-
porting filaments so conspicuous at later stages (Pl. I1.,
fig. 5, sup. fil.) The cell migrates towards the centre of
the tubule, becomes slightly elongated in a radial direc-
tion, and its outer end becomes frayed out into one or
more delicate filaments, which seem to retain some con-
nection with the wall of the tubule. The nucleus and
cell body then break down and disappear, and a group of
filaments is left arranged radially in the lumen of the
tubule, on which the spermatozoa formed at this time,
and subsequently are arranged in rows. In the smaller
and terminal portions of the tubules the whole epithelium
takes part in the formation of spermatozoa or ova, accord-
ing to the sex, but in the larger branches, and this is the
more common appearance in the sections, the tubule
possesses a longitudinal strip of columnar epithelium,
cells with clear cell bodies, and with long cilia projecting
into the lumen. ‘This strip of ciliated epithelium extends
round the tubule for about one-fourth or less of the cir-

SEA-FISHERIES LABORATORY. 115

cumference. It is more common in the male gonad, but
is present also in the female.

The parenchymatous tissue between the tubules has
much the same characters as in Stage I., but it 1s now
arranged in a more even system of bars and is not so
massive. Certain appearances, such as the difficulty of
distinguishing the limits of the growing tubules in relation
to the surrounding reticulum, seem to indicate that the
gonads grow at the expense of this tissue, or at least
absorb it during the process of growth. It was, however,
impossible to demonstrate this with certainty in the pre-
parations studied.

Ripening of the gonads proceeds slowly from now on to
the end of January, and consists of a gradual increase in
the width of the already-existing tubules and of the for-
mation of side twigs. The contents of the tubules slowly
increases, and it must be understood that from November
onwards there is an increasing number of free and
apparently mature ova and spermatozoa in the gonads.
The fact that moving spermatozoa may be obtained from
the mantle at the end of the year has been already referred
to. Whether the genital products present in the gonads
at the end of the year are fully mature and are capable of
development is uncertain, but if so, the presence of free-
swimming larve in the sea about this time may be easily
explained, since ova and spermatozoa may be swept out of
the lower partially ciliated tubules, and fertilization may
result. But a certain time may also be required for the
complete maturation of the genital products after their
proliferation from the germinal epithelium.

STaGe III. (Pl. IIL., figs. 1, 2)—About this time the
mantle lobes have attained the maximum of thickness, and
the gonads have invaded every part of the body un-
occupied by the other organs, even the outer wall of the

116 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

pericardial sac being overspread by them. The whole
space between the internal and external faces of the
mantle is now filled up by the gonads, and the tubules of
the latter are filled, in female and male, with a dense mass
of ova and spermatozoa respectively. The spermatozoa
are arranged in rows on the radial-supporting filaments
already mentioned, and the mutual pressure of the ova
against each other in slightly later stages than this gives
them a polyhedral form. The germinal epithelium is still
undergoing proliferation, but this is not so active now.
Where, in the male, the walls of the tubules are closely
compressed together, the basement membrane can be seen
as a thin, homogeneous line, on either side of which is an
irregular row of large, rounded cells, with clear cell bodies
and conspicuous nuclei, and internal to this the tubule is
filled with a mass of radially-arranged rows of spermatozoa.
The conspicuous reticular tissue present in earlier stages
is reduced to the merest trace, and is present mostly at
the angles formed at the junction of several tubules. It
stains less intensely with eosin, but the peculiar vacuo-
lated appearance already referred to still exists. Here
and there it is present as rounded masses possessing single
nuclei, but it is mostly fibrous.

- As in the male, the germinal epithelium still exists in
fic female, and ova are still being separated off. Most of
the eggs le freely in the cavity of the tubule in contact
with each other, but some are still attached to the wall by

a short stalk. In some parts the wall of the tubule seems
to consist of the basement membrane only.

STace IV. (Pl. IL., fig. 5).—From April onraba there
is apparently little change in the appearance of the gonads.
The maximum development, as indicated by the increase
in mass of the tubules, the numbers of ova and sperma-
tozoa. liberated from the. germinal epithelium, and. the

SHA-FISHERIES LABORATORY. Gy

reduction of the other tissues in the mantle, was obtained
in the specimens taken during April. Some taken later
were, indeed, less advanced, but generally this seems to be
the time at which proliferation from the walls of the
tubules mainly ceases. In July, Mussels were taken in
which spawning was in progress. In these the females
had almost completely spawned, and in the males the
gonads had undergone a great reduction in mass. A few
stray eggs lay scattered over the section situated in tubules
of greatly reduced diameter, one egg being present, as a
rule, in the cross section of the tubule, and generally
nearly filling it. Many tubules were completely empty.
The germinal epithelium persisted in some of these tubules
as one or more layers of small, rounded cells, but more
often its structure was difficult to make out.

The reticulum so apparent in Stage I. is already formed,
but in structure is more spongy than in the stage following
the complete extrusion of the ova, and the spaces between
the bars are about equal to those occupied by the tissue
itself. These interspaces have, In many cases, a cir-
cular outline, and probably represent the former situation
of ovarian tubules. The general histological characters of
the tissue are similar to those described in Stage I., except
that the vacuolated appearance is not so apparent.

The most striking changes have occurred in the male.
In some of the specimens taken at this time extrusion of
the spermatozoa has been almost completely effected, but
in others the animal had been taken in the act (PI. II.,
fig. 5). Here the sperm tubules have been greatly con-
tracted in cross area, and the mass of spermatozoa (PI. IL.,
fic. 5) within the tubule has become correspondingly
denser. The reticulum has increased enormously, and as
it seems difficult to derive this tissue from the exceedingly
fine fibrous network present in the last stage, it 1s possible

118 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

that it has spread into the spaces surrounding the tubules
of the gonad from other parts of the body. Round each
tubule this reticulum forms a dense mass, with few or no
interspaces, and its syncytial character is here strongly
suggested. The nuclei are not so evident as at a later
stage. Within the tubule itself the linear arrangement of
the spermatozoa is very striking, but instead of being
directed towards the centre these rows point towards that
part of the wall lined with the strip of ciliated epithelium
already described (PI. IT., fig. 5, ep. cil.). Often a blood
space (bl. sp.) is situated beneath this epithelium; over
the cilia and within the tubule is a space free from sper-
matozoa. The wall of the tubule has become indistinct
from the mass of spermatozoa which seem to abut directly
on the tissue of the reticulum (re¢.) surrounding the tubule.
The whole arrangement strikingly suggests the exercise of
pressure on the tubule by the rapidly developing reticulum
and the removal of the genital products brought into close
contact with the ciliated strips of the tubules. The lower
conducting portions of the gonads are entirely lined with
ciliated epithelium, and once here the genital products
must be speedily removed.

Briefly summarised then, the history of the yearly change
in the reproductive organs of the Mussel is as follows:—
(1) There is a short period of rest following spawning, and
occupying some part of August and September, during
which the space in the body of the animal formerly
occupied by the gonads is largely filled up by a large-celled
reticulum. This is followed by (2) the reformation of the
germinal epithelium and a slow proliferation of ova and
spermatozoa, lasting for the rest of the year. During this
period the gonads increase greatly in mass, and invade the
imantle and other parts of the body, displacing the reticu-
Juin formed during spawning. Succeeding this is a period

ee

SEA-FISHERIES LABORATORY. 119

(3) of active formation of the genital products, ending late
in the spring, after which ova and spermatozoa are formed
less rapidly. A slow emission of these now goes on until
in the summer, about June or July, the maximum spawn-
ing period of the Mussel occurs.

It is unlikely that any great proportion of the larve
found during the course of the year are spawned otherwise
than at the maximum spawning period in the summer
months. We have seen that it is probable that a certain
proportion are spawned during the first half of the year,
but that these probably result from the spawning of
isolated individuals or from the slow emission of ova and
spermatozoa prior to the act of spawning proper, and the
fact that this occurs does not materially affect the general
statement that the Mussel spawns in the summer. For
the year 1898 we have fixed this spawning period as
having occurred during July, but it is probable that it
is variable, and as the Committee’s bye-law fixes the
months of May, June, July, and August as those during
which it is illegal to take Mussels from the beds in the
district, it is very probable that, notwithstanding some
considerable variation in the occurrence of this period, it
will always fall within that close time. Probably spawning
is very generally over before August begins, but of this
further evidence is wanted.

EXXPLANATION OF THE PLATES.
PuateE I.

Fig. 1. Dissection of an adult Mussel to show the external
opening of the gonad. The right valve has been
removed, and the right gill and mantle lobe cut
away close to the junction with the visceral mass
and reflected upwards. Full size. «— indicates
the plane of the section shown in fig. 2.

120 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Fig. 2.

Bie sik:

Berea:

Fig. 38.

Hig. 4.

Complete transverse section through the body of
an adult female Mussel passing through the peri-
cardial cavity just posterior to the heart. Note the
presence of ovarian tubules in every part of the
section, the stage of development being the same
both in mantle lobes and in visceral mass. The
specimen was taken about the end of September,
and proliferation of ova from the germinal
epithelium is beginning, but the diameter of the
tubules is still very small. x 5 diameters.

PLATE II.

Transverse section of mantle of nearly ripe female
Mussel from Roosebeck outer scar, taken during
March. The mantle is nearly filled with the
tubules of the gonad containing many free ova.
The germinal epithelium is still undergoing
proliferation. xX 40 diameters.

Transverse section of mantle of nearly ripe male
Mussel from Roosebeck outer scar. (March.)
The mantle is filled with sperm tubules. x 40
diam.

Transverse section of mantle of female spent
Mussel from New Brighton beds, taken during
August. The tubules of the gonads have in many
cases broken down. T'wo are shown in the section,
one containing an egg which has not been
extruded during spawning. Note the great
development of the reticulum. X 200 diam.
Transverse section of mantle of spent female
Mussel from the beds in the Barrow Channel
(August). A blood space lying immediately
beneath the epidermis is shown, beneath which
are two collapsed ovarian tubules. Note that

SHA-FISHERIES LABORATORY. ca POF

each is lined partly by ciliated and partly by
germinal epithelium. X 200 diam. |

Fig. 5. Transverse section of mantle of spawning male
Mussel (New Brighton beds, July). A sperm
tubule is shown, partly lined with ciliated
epithelium. Note the great development of the
reticular tissue. > 230 diam.

Fig. 6. Transverse section of mantle of completely spent
female Mussel (Barrow Channel, August). xX 35
diam.

Fig. 7. Portion of transverse section of mantle of female
Mussel (Barrow Channel, September 30). A
developing ovarian tubule is shown. Actual
size=0'568mm. Note the proliferating germinal
epithelium, and the ciliated strip of wall. x 660
diam.

REFERENCE LETTERS.

abd., abdomen; add. a., anterior adductor muscle of
the shell; add. »., posterior adductor muscle of the shell ;
br. R., right gill; br. L., left gill; bl. sp., blood space; er.
st., crystalline style; ep. ger., germina] epithelium; ep.
cil., ciliated epithelium; ep. ext., external epidermis
of mantle; wzt., intestine; Mn. L., left mantle lobe; Mn.
f., right mantle lobe; per., pericardial cavity; per. gid.,
pericardial gland and auricle; pp. gen., right genital
papilla; pa. d., right dorsal labial palp; pa. v., right ventral
labial palp; ven., renal organ; ret., reticulum of mantle;
ret. bys., retractor muscles of the byssus; ret. bys. in.,
insertion of retractor of the byssus into the shell; rect.,
rectum; sup. fil., supporting filaments in sperm tubule;
st., tubular portion of the stomach; sp., rows of mature
spermatozoa; twb. ov., ovarian tubule; visc. nv., visceral
nerve cord.

122 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

On SEA-FisH HATCHING. ©
(W. A. HERDMAN.)

In the Twelfth Annual Report of the Sea-Fisheries
Inspectors for England and Wales (1898), under the mar-
ginal heading ‘“‘ Sea-Fish Hatcheries” (p. 29), Mr. C. E.
Fryer enters upon such a severe and detailed criticism
of the results attained that it cannot be regarded as
other than an attack upon the practice of artificial
hatching of sea-fish eggs as developed in America. It is
fair to say that it is the results of the hatcheries in America
that are attacked, as, although Scotland and Norway are
mentioned, these countries are treated very briefly, and no
conclusions are drawn from the few statistics quoted. The
greater part of the discussion deals, then, with Canada,
Newfoundland, and especially the United States. I gather
that Mr. Fryer has not himself visited the hatcheries he
discusses, and that his information is entirely derived from
the published annual reports—an unexceptional source, so
far as it goes, but one which may usefully be supplemented
by personal impressions and conversation with those
engaged in the work.

I regard Mr. Fryer’s paragraphs as a most useful state-

ment, drawing attention to the present position of the

hatching question from the critic’s point of view, counter-
acting any exaggerated opinions that may have been
expressed, and impressions that may have been enter-
tained, in regard to the immediate and enormous results
to be obtained from artificial operations, but requiring
some criticism, some modification, and some additions
before it can be accepted as a wholly satisfactory state-
ment of the matter. For although Mr. Fryer seems rather
carefully to avoid drawing any formal conclusions, he

SEA-FISHERIES LABORATORY. 123

certainly, consciously or unconsciously, by his manipula-
tion of the statistics, conveys to the reader an impression
entirely adverse to sea-fish hatching. And this impression
is, I believe, derived mainly from statements in regard to
the difficulty and not in regard to the efficacy of the
operations.

Mr. Fryer’s remarks are largely directed towards
removing impressions that may prevail as to the sim-
plicity of artificial hatching; and here we, in common
with all who have taken any part in such work, readily
concur in the general conclusion that the work is suffi-
ciently difficult and troublesome, if pains be taken to
attain success. I do not think that amongst those in this
country who have had to obtain in mid-winter or early
spring the spawning fish or the fertilized spawn, who have
had to filter the sea-water and watch day and night
its purity, salinity and temperature, who have jealously
guarded the embryos from danger and have tried to obtain
suitable foods for the larva, there can ever have been any
fond delusions as to the simplicity of the business. The
work is hard, there are many difficulties, the operation is
a delicate one, requiring constant and intelligent attention;
but all that need not deter. Given suitable conditions and
the right men, and the work can be successfully carried out.

While agreeing with Mr. Fryer that ‘“‘ simplicity” is a
delusion, I must demur against an impression which his
pages are lable to convey. It must be remembered that
the harrowing descriptions on pp. 36 and 37 of fish perish-
ing in the nets owing to the heavy storms, and freezing in
the boats and on the cars, and of embryos killed by the
million in the hatching boxes on account of the intense
cold, refer to conditions on the coast of North America,
and do not apply to our seas. These exceptional difficulties
either do not exist or are only present in quite a minor

124 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

degree for very short periods in our warmer and more pro-
tected coastal waters.

Turning now to the question of efficacy, we meet with this
difficulty—Mr. Fryer apparently does not accept the turning
out of millions of healthy young fry into the sea as a proof
of successful hatching operations. He seems to demand
a demonstration that the number of adult fish in the neigh-
bourhood has increased, without taking into account the
fishing operations that may be going on. In asking for
immediate proof that the fry live to become adult fish the
opponents of fish hatching are taking up what may seem
a very secure, but is, I consider, a very unreasonable position.
So many factors enter into the case that it is almost impos-
sible to devise a crucial experiment, or give immediate
scientific proof of a result. The demand may possibly be
satisfied in any locality next year, or the answer may be
delayed for 10 years or for longer; and yet the fisheries may
all the time be largely benefitted by—may owe their con-
tinued existence to—the artificially hatched fry added to
the population of the sea. And this may be the case even
if not a single individual out of the millions of fry set free
ever lies to become adult.

I do not think it at all probable that such unfavour-
able conditions ever exist, but I wish to point out that
even in such an extreme case the expenditure on land
and the sacrifice of life of the fry in the sea will not
have been in vain. If we try to realise the struggle
for existence in the sea, the toll that is taken at every
period of life from the egg to the adult, but which is excep-
tionally heavy in young stages, a moment’s reflection will
show that the addition in any natural hatching area of
some millions of young fish must distribute the danger of
being caught by an enemy over a much larger number
of individuals and so give to each a greater chance of

SHA-FISHERIES LABORATORY. °° -—s 125

escape. And if—as may be the case, we do not know—
the naturally hatched fry are hardier or more active, then
they will be the more likely to survive the perils, and will
owe their continued existence and their appearance in due
course somewhere as marketable fish to the presence for a
time in the sea of their brethren from the hatchery.

I shall come now from these general considerations—
to which Mr. Fryer does not seem to give enough atten-
tion—to some of the detailed statements in his report as
to the results of hatching, and upon some aspects of
which, such as the mortality in captivity, in my opinion
he lays far too much stress. I cannot now go into every
point upon which we may differ more or less, but shall
take up his treatment of the United States statistics, as
they are, perhaps, the most important, and the ones
discussed at greatest length.

After quoting (p. 39) the statistics of the Cod operations
at the Wood’s Holl Hatchery, he admits that the figures
show ‘“‘successful results.” He then attempts to upset
this conclusion by arguing that in computing the number
of eggs and fry derived from a parent fish we should take
into account not merely the actual parents, the fish that
spawned, but also all the others that were penned up in
the spawning pond. ‘This is surely a most unjustifiable
procedure. What reason is there to think that these
non-spawning fish would have spawned if left in the sea?
In fact, if the object is to make a comparison between
natural and artificial conditions, surely from what we
know of life in the sea it is highly probable that some of
the fish that spawned in captivity would, if left at large,
have been destroyed by man, or their natural enemies,
before they became parents. Consequently, I consider
that the Superintendent of the Hatchery was perfectly
right in considering as parent fish only those from which

126 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

spawn was obtained, and I should remove from Mr.
Fryer’s table, at the foot of p. 39, the three columns
headed ‘‘penned up,” as giving no additional useful
information.

It is also unfair to compare an average obtained by
including all the fish penned up, many of which did not
breed, with the total number of ova calculated as being
present in one selected large breeder. The comparison
can only be fairly made with the eggs in a similar number
of Cod, of the same size as those in captivity, chosen at
random from the sea. Even in that case the comparison
would only indicate whether there was a diminution in
the amount of spawn produced per captive fish. There
probably is such a diminution ; but, on the other hand, it
is certain that a large number of possible spawners in the
sea are killed by man and other enemies before or during
the spawning season.

Then, again, in making a further comparison between
artificial hatching and the ‘‘ processes of nature,”’ it is not
right to substitute (p. 41) the number of eggs calculated
to be present in the body of a parent for the actual
number of eggs treated in the hatchery. This method of
‘counting the chickens’’ not only before they are hatched
but even before the eggs are laid, is an ingenious method
of making the proportion of fry produced in the hatchery
seem very small; but I think the Superintendent’s figure,
54°6 per cent. of fry to ova, will commend itself to most
people as the correct calculation on the common-sense
basis of so many ova supplied and so many fry produced.
What would have happened to, say, 100 of the hatchery
spawners if they had been left in the sea, no one can say.
How many of them would live to spawn and how much
spawn they would produce we do not know. The per-
centage of fry to parent fish in the hatchery may, under

SEA-FISHERIES LABORATORY. 127

some circumstances, be small, but it may be smaller still in
a state of nature.

Té is curious how Mr. Fryer entirely ignores that side
of the question—the destruction in nature. He makes
a great point of the mortality in artificial operations,
and virtually assumes that in the sea every egg in
every fish will be spawned and will then hatch out.
This is entirely contrary to the experience of naturalists.
In addition to the destruction of the parents in the sea,
and the risks of non-fertilisation, and of being cast ashore,
we find at the spawning season nearly everything that has
a mouth (such as Meduse, Sagitta, Copepoda, and many
fishes) feeding upon fish eggs. One of the strongest
arguments in favour of the hatchery is that it protects the
embryos from their natural enemies. Judging from the
apparently stationary condition of the fish population in
the sea, from the enormous numbers of eggs produced, and
from our observations on the contents of fishes’ stomachs,
it is obvious that there must be a very great destruction of
eggs, and embryos and larve under natural conditions—
probably much greater than anything known in artificial
operations.

Moreover, in talking (as many do) of the small scale of
man’s hatching work compared with what takes place in
the sea, we do not sufficiently realise the meaning of what
is called ‘‘the balance of nature.”’ Millions of millions
of young are produced and these same millions of millions
are destroyed by natural causes, with the result that the
‘species remains fairly constant. If now man disturbs the
balance by catching some thousands of the adult fish in
a district, he is only equalizing matters and endeavouring
to minimise his destructive effect when he adds to that
sea-area some millions of artificially hatched fry. And
(theoretically at least—probably practically also, for correct:

128 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

theory and perfect practice must be in accord) he is the
better equalizer of the disturbed balance when he uses for
the hatchery the eggs of fish which would not have
spawned in a state of nature—that he can do by obtaining
ripe spawn from the fish trawled for market at the spawn-
ing season. This plan has been adopted in some hatcheries;
and in our own experiments at Port Erin and at the Piel
hatchery. It is the least expensive method of getting

spawn, but it is open to several difficulties and objections.

The other method is to collect the parent fish beforehand
and keep them in ponds until they spawn. It is the
method adopted in Norway and to a large extent in the
United States, and which Mr. Fryer has tried to show
results in a comparatively small number of fry per fish
used.

I shall not pursue the matter further. There is no need
for me to defend or expound the methods and statistics of
the United States Fish Commission. Their own officials
are very well able to answer for themselves, and very
probably they will do so—if they think it worth while.*

I have merely gone into some of the questions raised in
Mr. Fryer’s article because it seemed to me that he was
modifying the statistics in an unjustifiable manner, and
drawing conclusions with which I could not agree, and
which might have some bearing upon our procedure in this

* It may be well before leaving this part of the subject to point out that in
the appendix (a Manual of Fish-Culture, based on the methods of the United
States Commission of Fish and Fisheries, Washington, 1898) to their last
published report, which appeared only a few months ago, the United States
authorities remark in regard to their artificial propagation of the Cod:—‘‘The
unmistakable economic results which have attended these efforts warrant all
the time and money devoted to them and justify the greatest possible
expansion of the work.” I have also more recent letters from the Com-
missioner and the Naturalists on his Staff making detailed statements to the
same effect.

‘
4

SEA-FISHERIES LABORATORY. 129

country. We are not, however, in our work here so much
concerned with the actual details and difficulties of the
American work which Mr. Fryer criticises, as with those
general biological principles which I have emphasised
above—such as the destruction of young under natural
conditions in the sea, and the duty of man if he disturbs
the balance of nature in a locality to do what he can to
equalize matters by helping in the production and protec-
tion of the young.

With Mr. Fryer’s remark (foot of p. 41) as to breeding
from the larger and more vigorous parents I am in cordial
agreement. If aquiculture has been long delayed as a
scientific industry, it has as a consequence this advantage,
that it can adopt at once principles and practices evolved
gradually in the long history of agriculture and stock-
raising. It labours, however, under the obvious disadvan-
tage that so much of the result of our labour passes at
once beyond our ken. We cannot control the fish through-
out their life-history, we cannot yet take stock of the
population of our seas. This introduces such an element
of uncertainty into the problem, that, although I think we
have good reason to be encouraged and to continue the
work vigorously in a hopeful spirit and with an open
mind, still I for one would not go so far as to say (as some
have done) that marine fish-hatching had passed beyond
the experimental stage; and I may refer in this connection
to the somewhat fuller statement on the subject I made in
last year’s report,* and to my discussion there of the con-
ditions of a crucial experiment for the purpose of testing
the results of adding artificially hatched fry to the popula-
tion of a circumscribed sea-area. We await with interest
the result of the Fishery Board for Scotland’s work in

Loch Fyne.
* Report for 1897, pp, 24 and 25,

130 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

I cannot do better than conclude this section by quoting
a couple of sentences from a letter received during the
year from Professor M‘Intosh, whom we regard as the
pioneer of scientific fisheries work in this country. In
speaking of marine hatcheries, he says:—‘“‘ Of course such
institutions are strictly experimental, and it may be some
time before a decisive result is evident. Meanwhile, work
them thoroughly and support them liberally.’”’ I desire to
endorse that opinion. |

OYSTERS AND DISEASE.
(W. A. HERDMAN.)

In collaboration with my colleagues, Professor Boyce
(Bacteriologist) and Dr. C. Kohn (Chemist), I have been
working at this subject in our laboratories at University
College, Liverpool, for the last three years, and interim
reports upon the progress of the work have appeared in
our three last Fisheries Laboratory Reports. A detailed
paper, giving the full results of the investigation, has
lately been laid before the Royal Society of London, and
the main conclusions of that paper are as follows* :—

“1. Although our primary object was to study the
Oyster under unhealthy conditions, in order to elucidate
its supposed connection with infective disease, we found
it necessary to study in minute detail the histology of
certain parts of the body, especially the gills and mantle
lobes, the alimentary canal and liver. We give figures
and descriptions of these structures in both normal and
abnormal conditions.

“9, We have also worked out the distribution and
probable function of a minute muscle, which we believe

* Quoted from Nature for January 26th, 1899, p. 305.

SHA-FISHERIES LABORATORY. 131

to be the modified representative of the protractor pedis
muscle of some other molluscs.

«3. A diseased condition we found in certain American
Oysters very soon brought us into contact with the vexed
question of the ‘ greening’ of Oysters, and one of the first
results we arrived at was that there are several distinct
kinds of greenness in Oysters. Some of them, such as
the green Marennes Oysters, and those of some rivers on
the Essex coast, are healthy; while others, such as some
Falmouth Oysters, containing copper, and some American
Oysters re-bedded on our coast, and which have the pale
ereen ‘leucocytosis’ described in our former paper to the
Royal Society, are not in a healthy state.

‘““4. Some forms of greenness (e.g., the leucocytosis)
are certainly associated with the presence of a greatly
increased amount of copper in the Oyster, while other
forms of greenness (e¢.g., that of the Marennes Oysters)
have no connection with copper, but depend upon the
presence of a special pigment, ‘ marennin.’

‘““We are able, in the main, to support Prof. Lankester
in his observations on Marennes Oysters; but we regard
the wandering amoeboid granular cells on the surface of
the gills as leucocytes which have escaped from the blood
spaces, and have probably assumed a phagocytic function.

‘““5. We see no reason to think that any iron which
may be associated with the marennin in the gills, &c., is
taken in through the surface epithelium of the gill and
palps, but regard it, like the rest of the iron in the body,
as a product of ordinary digestion and absorption in the
alimentary canal and liver.

‘6. We do not find that there is any excessive amount
of iron in the green Marennes Oyster compared with the
colourless Oyster, nor do the green parts (gills, palp, &c.)
of the Marennes Oyster contain either absolutely or

132 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

relatively to the colourless parts (mantle, &c.) more iron
than colourless Oysters. We therefore conclude that
there is no connection between the green colour of the
‘Huitres de Marennes’ and the iron they may contain.

“7. On the other hand, we do find by quantitative
analysis that there is more copper in the green American
Oyster than in the colourless one; and more pro-
portionately in the greener parts than in those that are
less green. We therefore conclude that their green colour
is due to copper. We also find a greater quantity of iron
in those green American Oysters than in the colourless 3
but this excess is, proportionately, considerably less than
that of the copper.

‘8. In the Falmouth Oysters, containing an excessive
amount of copper, we find that much of the copper is
certainly mechanically attached to the surface of the body,
and is in a form insoluble in water, probably as a basic
carbonate. In addition to this, however, the Falmouth
Oyster may contain a much larger amount of copper in
its tissues than does the normal colourless Oyster. In
these Falmouth Oysters the cause of the green colour may
be the same as in the green American Oyster.

“9. By treating sections of diseased American Oysters
under the microscope with potassium ferrocyanide and
various other reagents,.we find that the copper reactions
correspond in distribution with the green coloration; and
we find, moreover, from these micro-chemical observations
that the copper is situated in the blood-cells or leucocytes,
which are greatly increased in number. This condition
may be described as a green leucocytosis, in which copper
in notable amount is stored up in the leucocytes.

“10. We find that an aqueous solution of pure hema-
toxylin is an extremely delicate test for copper, Just as
Macallum found it to be for iron,

SEA-FISHERIES LABORATORY. 188

“11, Experiments in feeding Oysters with weak solu-
tions of various copper and iron salts gave no definite
results, certainly no clear evidence of any absorption of
the metals accompanied by ‘ greening.’
~ 12. Although we did not find the Bacillus typhosus
in any Oysters obtained from the sea or from the markets,
yet in our experimental Oysters inoculated with typhoid
we were able to recover the organism from the body of
the Oyster up to the tenth day. We show that the typhoid
bacillus does not increase in the body or in the tissues of
the Oyster, and our figures indicate that the bacilli ponen
in the intestine.
~ “13. Our experiments showed that sea-water was
inimical to the growth of the typhoid bacilli. Although
their presence was demonstrated in one case on the
twenty-first day after addition to the water, still there
appeared to be no initial or subsequent multiplication of
the bacilli. |

“14. In our experiments in washing infected Oysters
in a stream of clean sea-water the results were definite
and uniform; there was a great diminution or total dis-
appearance of the typhoid bacilli in from one to seven
days.

“15. The colon group of bacilli is frequently found in
shell-fish as sold in towns, and especially in the Oyster ;
but we have no evidence that it occurs in mollusca living
In pure sea-water. The natural inference that the presence
of the colon bacillus invariably indicates sewage con-
tamination must, however, not be considered Eevabuiehed
without further investigation.

“16. The colon group may be separated into two
divisions: (1) those giving the typical reactions of the
colon bacillus, and (2) those giving corresponding negative
reactions, and so approaching the typhoid type; but in

184 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,

no case was an organism giving all the reactions of the
B. typhosus isolated. It ought to be remembered, how-
ever, that our samples of Oysters, although of various
kinds and from different sources, were in no case, so far
as we are aware, derived from a bed known to be con-
taminated or suspected of typhoid.

‘17. We have shown also the frequent occurrence, in
various shell-fish from the shops, of anaérobic spore-
bearing bacilli giving the characteristics of the B.
enteritidis sporogenes recently described by Klein.

“18. Consequently, as the result of our investigations,
and the consideration of much evidence, both from the ©
Oyster-growers’ and the public health officers’ point of
view, we beg to recommend—

(a) That the necessary steps should be taken to
induce the Oyster trade to remove any possible
suspicion of sewage contamination from the beds and
layings from which Oysters are supplied to the
market. This could obviously be effected in one of
two ways, either (1) by restrictive legislation and the
licensing of beds only after due inspection by the
officials of a Government Department, or (2) by the
formation of an association amongst the Oyster-
erowers and dealers themselves, which should provide
for the due periodic examination of the grounds,
stores, and stock by independent properly qualified
inspectors. Scientific assistance and advice given
by such independent inspectors would go far to
improve the condition of the Oyster beds and layings,
to re-assure the public, and to elevate the Oyster
industry to the important position which it should
occupy. |

‘“(b) Oysters imported from abroad (Holland,
France, or America) should be consigned to a member

SEA-FISHERIES LABURATORY. 185

of the ‘Oyster Association,’ who should be compelled by
the regulations to have his foreign Oysters as carefully
inspected and certified as those from his home layings.
A large proportion of the imported Oysters are,
however, deposited in our waters for such a period
before going to market that the fact of their having
originally come from abroad may be ignored. If this
period of quarantine were imposed upon all foreign
Oysters a great part of the difficulty as to inspection
and certification would be removed.

“‘(c) The grounds from which Mussels, Cockles,
Periwinkles are gathered should be periodically
examined by scientific inspectors in the same manner
as the Oyster beds. The duty of providing for this
inspection might well, we suggest, be assumed by the

various Sea-Fisheries Committees around the coast.”’

Norse ON OCCURRENCE OF IRON AND COPPER IN OYSTERS.*
By CHartes A. Koun, B.Sc., Ph.D.

‘The investigations of Professors Herdman and Boyce
on the life conditions of Oysters, which have been in
progress since 1895, have pointed to the desirability of
ascertaining the quantities of iron and of copper they may
contain under either normal or abnormal conditions.

‘Two points of interest have arisen in this connection.
In the first place the relation of iron to the greenness of
the healthy French Oyster (Huitre de Marennes); and
secondly, the extent to which copper is responsible for the
pale green colour of American and other Oysters, a
diseased condition accompanied by a leucocytosis dis-
covered and especially studied by Herdman and Boyce.
Lhe presence of winute quautitics of copper and of iron

* Quote] from Report of Oyster Committee to Brit. Assoc., 1898.

136 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,

as normal constituents of all Oysters has been shown by
the analytical data obtained.

‘The results recorded have been made at Professor
Herdman’s request, and have proceeded side by side
with his investigations. Now that these are completed,
a summary of the work from, a more purely chemical
standpoint may be of interest, especially since the occur-
rence of these metals—copper and iron—either from the
point of view of the origin of colouration or the cause of
poisoning has from time to time been the subject of
discussion.

‘The Analytical Method Employed.—Electrolytic
methods of analysis were adopted both for the determin-
ation of iron and copper: these methods, I have already
shown,* possess marked advantages for the estimation of
minute quantities of metal, especially if derived from
organic matter, for they are quite free from any prejudicial
influences traces of organic matter may exert, such
as arise when volumetric or colorimetric methods are
employed. In each determination the bodies or gills only
of six or more Oysters were carefully washed, dried
between filter paper to remove as much adherent moisture
as possible, and then carefully dried in porcelain dishes in
the air bath at 100° C. When this drying was as com-
plete as possible, the Oysters were heated in the air bath
until thoroughly carbonised, the carbon carefully burnt off
over the free flame, and the residue finally ignited in a
porcelain crucible. Special care was taken to exclude
dust during both the drying and the ignition. The ash
was then thoroughly extracted with a mixture of 25 c.c.
hydrochloric acid and 25 c.c. sulphuric acid (1:2) on the
water bath, and the resulting solution filtered and

‘‘ Brit, Assoc. Rep., 1893, p. 726.

SHA-FISHERIES LABORATORY. 137

concentrated. The residue was free from both copper
and iron. The acid solution obtained was electrolysed
for copper with the usual precautions, a spiral of fine
platinum wire weighing about 5 grme. being employed as
the cathode. The iron was determined in the residual
solution, after neutralisation with ammonium hydrate, &c.,
acidifying with a few drops of oxalic acid solution, and
boiling with ammonium oxalate: 4 grme. of the oxalate
were added in each case, the precipitated calcium oxalate
(which is quite free from iron) filtered off and thoroughly
washed and the resulting solution electrolysed, the metallic
iron being also deposited on a spiral of platinum wire. A
blank experiment with all the reagents employed was
made, and the amount of metal found (0°002 grme. iron)
deducted in each case. Also the deposited metal, both
iron and copper, was dissolved off the electrode by acid,
the solution obtained tested by the ordinary reagents and
the spiral re-weighed, as a check upon the determinations,
since the quantities found were extremely small.

‘The Green Colour of French Oysters, ‘‘ Huitres de
Marennes,” and the Presence of Iron in Oysters.—The early
observations of Dumas (1841) and of Berthelot (1855)
showed that the green colour of ‘‘ Huitres de Marennes”’ is
not due to chlorophyll, and that although every Oyster
contains a certain very small amount of copper in its
blood in the form of ‘“‘ hemocyanin,” as determined by
Fredericq, the green colour of the French cultivated
Oyster is not due to this metal. Ray Lankester* in 1886
confirmed the latter statement, and states in his investi-
gation on the histological condition of the colour that
there is neither copper nor iron in the refractory blue
pigment ‘‘marennin”’ of the coloured portions of the

* Quart. Journ. Micros. Sci., 1886, 26, 71.

138 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Oyster. Berthelot, however, suggested that the green
colour was due to iron, and more recently Chatin and
Muntzt have extended and corroborated this statement.
- From their analytical results these observers conclude
that both the green and the brown colourations of various
types of French Oysters are due to the presence of
iron, and that the depth of colour bears a close pro-
portion to the quantity of iron contained. The coloura-
tions are chiefly apparent in the gills, but extend also to
the labial palps and parts of the alimentary canal. Chatin
and Muntz base their conclusions in the first place upon
the fact that they find considerably more iron in the gills
than the rest of the body of green Oysters; and secondly,
upon the occurrence of a larger quantity of iron in the
eills of green than of white Oysters.
‘Appended are some of their results, to which I have
added a column, showing the ratio of the iron in the body,
minus gills, to that contained in the gills.

Iron per 100 parts of

dried organic matter.
Oyster. Colour. sae a
UY apa
1. Gils)
*| of body
Cancale... ...| White wan ...| 0°03879 0°0241 1°57
Areachon ... seo RNS een oe ...| 0°0605 0-0357 1°69
Marennes ... nan| Ely Seen re seat OSO702 0°0318 1°21
Cancale... ...| Brown green ...{ 0°0804 0°0476 1°69

Sables d’Olonnes ... Very dark brown | 0:0833 0°0436 ion
green

- ©The relative proportion of iron in the gills hardly bears
out the conclusions arrived at; it is the same in pale-
green and brown-green Oysters, and in both, but little
greater than in the white. On the other hand, the total

+ Compt. Rend., 1892, 118, 17 and 56,

SEA-FISHERIES LABORATORY. 139

iron, both in the gills and in the rest of the body,
shows a marked increase, apparently corresponding to the
depth of the colouration. The iron was determined in
these experiments by potassium permanganate, but the
absolute quantities of metal found are not stated. The
calculation of the results per 100 parts of dried organic
matter is apt to be misleading. In my own experiments
it was not found possible to get anything approaching
constant weights in this way, and the results are entirely
out of accord with those of Chatin and Muntz.

‘The following table gives the quantities of iron found in
French as compared with American Oysters, three pairs
of gills being analysed in each case.

‘These figures show conclusively that there is more and
not less iron in the gills of the white American Oysters
than in the French, and this irrespective of the basis on
which the result is calculated. The ash is undoubtedly
the most reliable factor to calculate on, provided the
Oysters are carefully washed before drying, which was
always done: the result per pair of gills (or Oyster) 1s
most in accord with this, and has the advantage of being
an easy and in many respects useful basis.

Huitres
— de American.
Marennes.

Gross body weight, after dr nying between

filter paper . 3°8 grms. | 6°5 grms
Weight dried at 100° C. "for six hours: ealQnoe i. L025 ta.
Weight of ash ... O0940n. 0°1140 ,,
Weight of Zvon found . 0:00037 |; 0:0008 ,,
Ratio of Iron found :—
(1) Calculated per pair of gills 1 to 2°7
(2) 53 on gross body weight.. 1 to 1°56
(3) re at weight at 100° ey 1 to 1°36
(4) 9 on ash ... ; 1 to 2°2

140 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

‘The relative quantities of iron present in the gills as
compared with the rest of the body were next determined |
in French, Dutch, and American Oysters. Six Oysters,
or the gills of six Oysters, were analysed in each case
with the following results :—

Weight of Iron found in

Six Oysters. aad
French. | Dutch. | American.
Gillsie. Eis = ct as ie 0°6 0°4 2°3
Bodies minus gills ... ee sia 1°2 15 14
Weight of ash of gills... ahs -=.| O°1880 0°0217 0°0294
| as »» Of bodies minus gills ...| 0°5980 0°1125 0°1240
7% Irononash. Gills... oe Neeley) 1°85 7°82
aa “i Bodies minus gills... 0:20 1°33 1°37
Ratio of Iron in gills to Iron in rest of
body.
Calculated per Oyster 55 52\ ee? 1\23545 1:0°74
An Olash 2. ox op | 1G a 1: 4°) 7 Dye

‘From these figures it 1s evident that the gills of the
French Oysters do not contain an excessive quantity of
iron such as might account for their colour. Calculated
per Oyster the gills contain less iron than the rest of the
body, except in the American Oyster; calculated as a
percentage on the asl the reverse is the case. The |
proportionate quantity of iron in the gills as compared
with the rest of the body is somewhat greater in the
French Oysters than in the Dutch, but much less than in
the American.

‘Clearly, therefore, there is no connection between the
green colour and the quantity of iron present. This result
igs quite in accord with Ray Lankester’s observation that
his ‘‘ marennin”’ is free from iron as well as from copper.

‘Both the gills and bodies of Oysters contain a small —

SEA-FISHERIES LABORATORY. 141

quantity of iron, which is evidently normally present, the
gills containing a somewhat larger amount in proportion
to the total quantity of mineral matter present.

‘Finally, the total iron in a variety of Oysters was
determined in order to ascertain the normal quantity
present. ‘These data, which are tabulated below, show a
fairly constant proportion of iron per Oyster, from 0°15 to
0°36 mgrme., or from 0°18 to 0°65 per cent. on the ash.

Total Iron present in Oysters.

| Megrme. | Percent-

; Number |Total Iron| Weightof| Iron age of

Variety of Oyster | Analysed.| grme. Ash. per Iron on
Oyster. Ash.

*“Huitres de Mar-

merrennes ... 6 0°0018 0°7860 0°30 0°23
Dutch 6 0:0009 0°1393 0°15 0°65
American .., 5 0:0018 0°2791 0°36 0°64
Colne 10 0:0020 1°0938 0°20 0°18
Deep Sea .... 2 0°0064 15017 0°32 0:43
Falmouth ... 6 0°0016 0°4534 | 027 0°35

‘In considering the variations in quantity, the very small
amounts of metal present must be borne in mind. | |

‘Tt may be added that although Carazzi has attributed
the green colour of French Oysters to iron taken up from
the mud of the Oyster-park or “claire,” experiments on
feeding Oysters with very dilute solutions of iron salts
(0:02 to 0°01 per cent.) carried on in conjunction with
Prof. Herdman produced no green colouration whatever.
The only result was a certain amount of ‘‘ browning”’
throughout the Oyster, the gills being no more affected
than the rest of the body. More recently Carazzi has
shown that Oysters fed with similar dilute iron solutions
acquire a pale yellowish colour in certain parts (branchial
epithelium and the cesophageal mucous membrane), and

142 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

that in these parts microscopic tests show the presence of
granules of iron. The actual meaning of these results
can hardly be recognised without quantitative data.

‘The Presence of Copper in Oysters.—F redericq has shown
that a certain small amount of copper is present normally
in the hemocyanin of the blood of crustaceans and
molluscs. The quantity thus present in Oysters of
different origin is fairly constant as shown in the following
table: it varies from 0°25 to 0°66 mgrme. per Oyster, or
from 0°30 to 1:18 per cent. on the ash.

|

Mgrme. | Percent-
tf Number | Total |weight of] Copper age
Variety of Oyster. | Analysed. Copper Ash. per | Copper on
Seles Oyster. Ash.
‘Huitres de Mar- |
rennes’ ... bee 6 0°0024 0°7860 0°40 0°30
Dutch 6 0°0015 0°1393 0°25 1°08
American 5 00033 0°2791 0 66 1°18
Colne 10 0:0036 1°0938 0°36 0°33
Deep Sea 2 00069 15017 0°34 0°46

‘0-4 mgrme. per oyster may be taken as an average, a
quantity slightly greater than the average iron (0°26
merme.). The calculated percentages on the ash show
greater variations, due to the very considerable differences
in the total quantities of mineral salts present, and it is
probably to this last factor that the popularly recorded
differences in taste of the various kinds of oysters is really
due. Certainly the minute quantities of copper and iron
present cannot account for them.

‘The copper was also determined in the gills and in the
bodies minus gills of French, Dutch, and American
Oysters, with the following results :—

SEA-FPISHERIES LABORATORY. - 143

— Determination of Copper.

French Dutch. American.

Six Oysters. ‘Huitres de Marennes.’

Trace

Gills only ... “ie ai
“ae -4 mgrme.

Bodies minus gills

=)
CO
co ~T

oo

‘These data show conclusively that the green colour of
the gills of French Oysters is also in no way connected
with the copper present.

‘Quantities of copper greater than those recorded point
to abnormal conditions. Such have been found to occur
with certain Falmouth Oysters, and in an especially
interesting manner with the green leucocytosis of American
and Falmouth Oysters—the diseased condition referred to
above.

‘Falmouth Oysters.—The presence of relatively large
quantities of copper in Falmouth and other Cornish
Oysters has been repeatedly associated with their bluish-
ereen colour. Dr. T. EH. Thorpe* states that these
Oysters, the colour of which, both in character and dis-
tribution, is quite. different from that of the Marennes
Oysters, contain on the average about 1:3 mgrme. of
copper per Oyster. ‘This large proportion is, Dr. Thorpe
says, ‘obviously caused by the mechanical retention of
cupriferous particles.’ On relaying they lose their

colour, and the quantity of copper present becomes

normal, 0°'4 mgrme. per Oyster.

‘Six Falmouth Oysters, the bodies of two of which were
of a distinct arsenic-green colour, were dried at 100° C.,
and then digested with water and subsequently with dilute
hydrochloric acid. The extract contained about half the
total copper present, showing that the metal is partially,

* “Nature,” 1896, p. 107.

144 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

at any rate, mechanically retained on, or in the body of
the Oyster, probably as a basic carbonate.
‘The analytical results were as follows :—

Mgrme. | Mgrme. | Per cent. | Per

f Cop- Weght.| Copper Iron Copper | cent.
Six Oysters. per. | Iron. Jof Ash. per per on. Iron on

Oyster. | Oyster. Ash. Ash.

Extract with
dilute acid ...| 0°0097| 0:0024| 0°2272| 1:62 0°40 4:22 1:06
Oysters ...| 0°0114} 0:0016| 0°4534| 1°90 0:27 2°51 0°35
Total ...| 0'0211! 0°0040} 0°6806| 3°52 0°67 3°10 0°59

|

‘The total copper present is almost nine times the
normal quantity, and about half of this is easily removed
by dilute acid. It is quite likely that the remainder is
partially or wholly simply entangled in the food passages
of the Oyster, and that the green colour may be due to
some other cause than this mechanically retained copper,
as suggested by Herdman.* Mr. G. C. Bourne, indeed,
regards it as due, in some Falmouth Oysters, to a green
desmid upon which the Oysters feed in quantity.

‘The occurrence of copper under such conditions is due
to the locality, and may quite possibly attain injurious
proportions, for the Oysters were obtained from a creek
which is locally supposed to bring down copper, and the
mud of which was found by Thorpe to contain 0°148 per
cent. of copper. Normal sea-water contains such an
excessively small quantity of copper that it was not found
possible to detect its presence, even electrolytically, in a
litre of sea-water, after concentration. .

‘The green leucocytosis already referred to was first
noticed by Herdman and Boyce in American Oysters

* * Nature,” 1897, p. 366.

—

SEA-FISHERIES LABORATORY, 145

which had been relaid near Fleetwood. The colour
manifests itself in patches and streaks of pale green on
the mantle, in engorgements of the blood vessels and in

masses of green coloured leucocytes in the heart. The

leucocytes are apparently all amoeboid wandering cells,
comparable to the colourless corpuscles of the blood of
higher animals, and the colouration coincides with their
distribution. | |

‘The six greenest and six whitest of 120 of these Oyster
were chosen for analysis; also a quantity of the greenest

portions of the greenest Oysters was selected from another

batch, and compared with the corresponding portions of
the whitest Oysters. The iron was not determined in the
latter comparison, owing to contamination of metal in
cutting.

‘The following were the results obtained :—

; Mgrme. | Mgrme. j Per cent. | Per
Cop- Copper Iron Copper | cent..
_ Oysters. per. | Iron. | Ash. per per on Iron on
Oyster. | Oyster. Ash. Ash. |
Green ...| 0°0158] 0°0091} 171450; 92°68 1:52 1°38 0°79
White ...| 0°0042| 0°0036] 1°0948; 0:70 0°60 - 0°38 0°33
Greenest parts| 0°0033| — | 0°0780 -- — 4:23 —_
Whitest parts} 0°0009) — |0°0452 es | = 1°99 eas
|

‘The excessive quantity of copper in the selected green
Oysters is 3°75 times that in the white calculated per
Oyster, and 3°63 times calculated on the ash. In the
selected parts the total copper present calculated on the
ash is high in both cases, and the green parts again show
a marked excess in the proportion of 21 to 1. The copper
and iron in the white specimens are about normal, but
the increased quantity of iron in the green is marked,
being 2°5 times that of the former. Still there is relatively

146 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

a large excess of copper as compared with iron in the
green Oyster, as is evident from the ‘analyses, the ratio
being 1'1: 1 for the white and 1'8 : 1 for the green,

‘It is to be concluded, therefore, that the green colour —
of these Oysters is coincident with the distribution of the
excessive quantity of copper present, and that the copper
is in consequence to be regarded asthe cause of the
colour. The histo-chemical investigations of Boyce and
Herdman have amply confirmed this conclusion. |

‘Further, this leucocytosis is not accompanied by a mere
redistribution of copper, but by an absolute increase of
the amount present in the body. |

‘The deposition of copper in this manner is regar ded by
Boyce and Herdman “‘ as a degenerative reaction, due to
a disturbed metabolism, whereby the normal copper of
the hemocyanin, which is probably passing through the
body in minute amounts, ceases to be removed, and —
so becomes stored up in certain cells.” The change
is comparable in kind to the accumulation of iron in
pernicious anemia.

‘The increased quantity of iron present may also be due
to abnormal conditions of life, but a more accurate
localisation of the normal iron of the Oyster is necessary
before this can be decided. as.

‘This green leucocytosis has been observed by Herdman
and Boyce in other Oysters, including those of Falmouth,
and it is likely to be the real cause of their colour; a
colour therefore due to copper as previously supposed,
but accompanied by a diseased condition. Whether the
presence of copper in the water facilitates in any way the
development of the disease has not been determined ;
experiments made on keeping Oysters in very dilute saline
copper solutions give no affirmative results beyond a
certain amount of post-mortem green staining.

4
-
a
4
.
..
-

)

;

SEA-FISHERIES LABORATORY. 147

‘Manganese was found to be present in several of the
varieties of Oysters analysed. Its detection is readily
effected in the electrolytic method of analysis as it
separates at the anode as peroxide. Colne Oysters con-
tained 0:14 mgrme. per Oyster—a rather smaller quantity
than the iron found.’

MUSSEL-BEDS AND MUD-BANKS.
By hh. i. ASCROFT.

On the Lancashire and Cheshire coasts it is often
noticed that Mussel-beds are situated on banks of mud.
Mussels require, in the earliest shelled stage, some
hard substance to attach themselves to, such as stone,
gravel, hardened sand (such as Sabellaria tubes, for
instance), shells, such as Cockles or Mussels, wooden
piles, and the bottoms of boats. As they grow up and
increase in size (when located where there is little wave
action), through their excreta and the mud settling
amongst them they are inconvenienced, and to escape
being buried a lengthening of their cables, or byssus, takes
place, and so they lift themselves. 'This process, repeated
time after time, raises the Mussels a great height above
their original location, and the bed of mud increases
accordingly.

The greatest depth that I have heard of occurred in the
River Ribble, where a bed of gravel was bare in the
channel a little above Lytham, when a strike of Mussels
occurred on it, and in the course of three years the
Mussels accumulated a bank of mud ten feet in depth—
there being no wave action to affect it.

At St. Annes-on-the-Sea, below the pier, there 1s a bed
of gravel on which a strike of Mussels occurs every two
years. During the two years growth the mud gets to a

148 TRANSACTIONS LIVERPOOL BIOLOGICAL SCCIETY.

depth of two feet, and then the heavy seas reach the bed
‘and roll the Mussels off like a carpet, and I have seen
them strewn along the beach at high-water mark as far as
Lytham Pier, a distance of four miles, during the same
‘tide. ay! <a
Under the refreshment rooms at Southport Pier is a
foundation composed of stone’on which Mussels strike.
They then accumulate mud until their cables, or byssus,
reaching down to the stones, get too weak, and they
are then washed away. The same thing occurs in
the Rock Channel, and on the great scars at; Heysham.

In the case of St. Annes, and doubtless in the other
cases also, if the seaward edge of the scar was protected
by piles or posts, and the Mussels so rendered less liable
to be broken into by the sea, they would then, in all
probability, be able to grow to a marketable size.

The researches of Mons. Viallanes at Arcachon, in the
South of France, have proved the power of Mussels in
comparison with Oysters to separate food and refuse (sand,
&c.) from the water. By his experiments he shows that
for one quart of water filtered by a French or native
Oyster 18 months old, a Portuguese Oyster of the same
age filtered 53 quarts, and a middie-sized Mussel 3 quarts.
The excreta respectively were 0°199 gramme, 1°075
crammes, and 1°768 grammes per 24 hours. So that in
proportion to the numbers covering the same area of
ground the Mussel will deposit three times as much
material as a Portuguese, and eighteen times as much as
a French or native Oyster.

There is no doubt but that the rate of growth is to a
ereal extent proportional to the food consumed, so that it
is clear that the Mussel has a much greater quickness .
growth than a native Oyster.

The accumulation of mud (other conditions being the

ay oth oe: eee,

SHA-FISHERIES LABORATORY. _ ry 4g

same) is much eréater where brackish water flows at
times over the bed, and it is a well known fact that
Mussels in such a place thrive and grow much more
quickly than those in pure sea water, no doubt through
the large quantity of food brought down by the rivers.
So much is this the case that every year Mussels are
removed from the scars on the seaward side of the River
Wyre below Fleetwood and placed in the same river above
Fleetwood to have the benefit of the brackish water.

The number of young Mussels (eggs and embryos) given
off by their parents must be in quantity simply enormous.
At Southport during the autumn, if a rope’s end is
allowed to hang in the water for the space of a month
you find it at the end of that time coated with Mussels,
like a swarm of bees hanging on a bough.

Boats, buoys, and all wrecks or posts near low-water
mark are smothered with them. These objects form
natural bouchots from which the young Mussels might
well be removed for cultivation elsewhere.

REPORT ON THE PHYSIOLOGY OF COLOUR-CHANGE IN
HIPPOLYTE AND OTHER MARINE CRUSTACEA.

By F. W. Keesue, M.A., and F. W. GampBur, M.8c.,
Owens Coilege, Manchester.

By the recommendation of Prof. Herdman and the
permission of the Lancashire Sea-Fisheries Committee,
we were allowed to utilise the Piel Laboratory for the
purpose of investigating the colour-physiology of
Hippolyte varians. We paid two short visits (from May
27th to 30th, and July 1st to 5th) in order to ascertain
the resources of the Laboratory and to discover the

localities in which Hippolyte could be obtained. As these

150 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

visits were successful, it was decided to spend the long
vacation at Piel, and accordingly the work was started
on July 15th and carried on without interruption till
Sept. 22nd., concluding with a visit Dec. 15-22. In the
present Report we merely give a short account of the
methods employed, since the results obtained by their
means are not yet ready for publication.

Hippolyte varians is one of the few Crustacea which
may be considered abundant in the neighbourhood of
Piel. It keeps, for the most part, to beds of weeds below
low-water mark, and hence its habits have largely to be
learnt from specimens in captivity. Compared with its
associates—Idothea, Mysis, and Crangon—Huppolyte is of
a delicate constitution, and needs constant aération or
constant change of sea-water to maintain it in good health.
The resources of the tank-room at Piel enabled us to
overcome this difficulty. Fresh weed or the dead bodies
of its fellows serve Hippolyte as food. With due
precautions specimens may be kept under observation for
ten days or a fortnight.

From the nature of the beds of sea-weed in the Barrow
Channel a haul with weighted canvas tow-nets usually
contains an assortment of Hippolyte of different sizes and
colours. Shades of brown and yellow are abundant,
whilst green and red are sometimes common, sometimes
rare. With the large Halidrys siliquosa a dark brown
variety is associated; among the fine Polyzoon
(Bowerbankia) which clothes the lower parts of the
Halidrys stems, a speckled variety of Hippolyte occurs :
in the tide-pools of Foulney Island the green variety, and
it alone, 1s found among the Zostera.

The methods employed for investigating whether there
is a power of colour-change, and if so, how this power may
be tested, were for the most part quite simple. One

SEA-FISHERIES LABORATORY. 151

method consisted in testing the effect of light of differing
intensities upon a given variety. ‘To do this a number of
jars, some of clear glass, others enveloped with muslin,
others again with black cloth, were taken on board, and as
soon as the nets were hauled up, the Hzppolyte were
recorded, and specimens of each variety were placed
in each jar. The effects produced differed very markedly
according to the time of day at which the catch was made ;
the result of treating a haul in this way after dark or even
in waning light being very different from that obtained at
mid-day.

We also directed our attention to the influence of
variously coloured weeds, with the object of testing
whether a brown Hippolyte would become green when
placed with green weeds, and red when surrounded with
red ones. For this purpose we had devised an apparatus
which was described shortly before the British Association
at Bristol.

This instrument consists of a ‘‘ pressure-bottle,” hold-
ing a couple of gallons of sea-water, which is discharged at
a uniform rate through an escape-tube commencing with
a minute opening. The water is made to circulate
through a series of air-tight observation- dishes, in which
the Hippolyte ave placed. The loss of water from the
pressure-bottle is made good by the entrance of air which
is first drawn through a second series of observation-
dishes, the water in which is changed once or twice a day
only. On the whole it was found that Hippolyte flourishes
best in the air-circulating dishes.

The influence of monochromatic light was investigated
by the aid of Landolt’s fluid ‘‘ colour-filters.” In order
that the light which reaches Hippolyte may be of fairly
high intensity and not merely equivalent to shade, we
employed a mirror to reflect sunlight or the light of an

152 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

incandescent lamp into the jars which were otherwise
quite opaque. .

The effects of direct sunlight, reflected light and
scattered light upon the colour of Hippolyte were tested
by exposing carefully recorded specimens to these in-
fluences in the open grassy space outside the Laboratory.
To prevent a rise of temperature in the sea-water, we
employed a circulation of water derived from the supply
in the Laboratory to flow through our observation-dishes.
The methods we used for investigating the effects of
rise and fall of temperature, electrical stimulus and of
toxic agents do not call for remark here. At night and
often during the day the records (chiefly microscopical)
were made by the hight of an incandescent lamp.

We have arrived at the conclusion that there are two
colour-phases in Hippolyte varians; one diurnal, the
other nocturnal. The recurrence of these phases is to
some extent independent of the conditions of illumination,
although the colour itself may be profoundly influenced
by varying the quality and intensity of the incident light,
and also by other stimuli, which do not act through the
eye.

The advantages which the Piel Laboratory affords for
work of this kind are very considerable, and while express-
ing our thanks to Professor Herdman and to the Curator
of the Laboratory (Mr. Andrew Scott), we wish to
acknowledge our marked indebtedness to the Lancashire
Sea-Fisheries Committee for the generous help which is
afforded by them for the prosecution of scientific work,
generosity which will surely become more widely
appreciated as it becomes more generally known.

Ce

en

SEA-FISHERIES LABORATORY. 153

ADDENDUM.

The species of Crustacea we worked with, have been
kindly identified by A. O. Walker, Esq. From the very
limited fauna of Piel shore, it may be of interest to give
the list, which, however, is not quite complete.

Hippolyte varians, Leach, common, just below the
level of ordinary spring tides.

Hippolyte fascigera, Gosse, a doubtful species.
Almost certainly a variety of H. varians.

$e Less common but oc-
Hippolyte cranchu, Leach.

curring with the
Huppolyte pusiola, Kroyer.

foregoing.

A species of Mysis which gave interesting results and
which occurs with Hippolyte has been determined as
Mysis neglecta, G. O. Sars.

A small collection of Pycnogonida, which have no con-
nection with the present research but which were taken
among the fine red weeds of the Barrow Channel, contains
the following forms :—

Ammothea echinata, Hodge.
Anoplodactylus petiolatus (Kroyer).
Pallene brevirostris, Johnst.
Nymphon gracile, Leach.

APPENDIX.

LANCASHIRE SEA-FISHERIES HATCHERY
AND LABORATORY AT PIEL.

REGULATIONS IN REGARD TO WORKERS.

1°. Biologists and Students desiring to work at the
Piel Hatchery should apply to the Hon. Director
(Professor Herdman) who, if there is room, will allot
them work places in the Laboratory in the order of
application. |

2°. In the absence of the Director, the Resident
Assistant (Mr. Andrew Scott) will determine which
places in the Laboratory workers are to occupy, and to
what extent the instruments of the Laboratory (micro-
scopes, microtomes, &c.), and the boats and collecting
apparatus may be used by workers.

83°. The Aquaria in the tank house are intended for
experiments in Fish Hatching and Fish Rearing, and it
is oply by express permission of the Director or the
Resident Assistant that they may be used for private
investigations.

4°, Laboratory accommodation and lodging in the
house are given free of charge to those duly qualified
workers or students whose applications have been
accepted, and who have been assigned places in the
Laboratory. Meals (breakfast, dinner, tea and supper)
are provided for those working in the Hatchery at a fixed
charge of 8s. per day or 15s. per week, payable to Mr.
Scott.

a
” bes a

SEA-FISHERIES LABORATORY. 155

5°. The Resident Assistant will be ready to give
assistance to workers at the Hatchery, and to provide them
with material for their investigations so far as it does not
interfere with his routine duties and his ‘‘fisheries”’
work.

6°. All dishes, jars, bottles, tubes and other vessels in
the Laboratory may be used freely, but must not be
taken away from the Laboratory. If any workers desire
to make, preserve and take away collections of marine
animals and plants, they must provide their own bottles
and preservatives for the purpose.

7°. The fish and other specimens in the tank room
are the property of the Institution and must not be used
or disturbed by workers in the Laboratory.

8°. Hach worker in the Laboratory is required to send
a short account of his work done at the Institution, and
of the results he has attained, to the Director before the

ist of December (at latest), in order that it may be

entered in the Annual Report to the Sea-Fisheries
Committee.
W. A. HERDMAN,

Honorary Director of the Scientific Work
to the Lancashire Sea-Fisheries
Committee.

156

NOTE on MID-WINTER SURFACE and DEEP
TOW-NETTINGS in the IRISH SHA.

By Isaac C. THompson, F.L.S.
[Read March 17th, 1899.]

TAKING advantage of a fine day and smooth sea, after a
month of abnormally boisterous weather round our coasts,
on January 29th, 1899, I accompanied Mr. Dawson,
Superintendent of the Lancashire Fisheries, and Prof.
Herdman, on an inspection of the spawning grounds off
the west of the Isle of Man, in the fishery steamer ‘‘ John
Fell.” In addition to the apparatus and trawls used on
boarl the steamer, we were provided with dredges and
tow-nets belonging to the Port Erin Biological Station,
our point of departure.

Gradually steaming up the western side of the Island,
when several hauls of the tow-nets and dredge were made,
it was thought by our scientific expert to be an excellent
opportunity for noting, on a typical winter’s day, both
the quantity and comparative character of the plankton
collected, under similar conditions, at the surface and at
the sea bottom. ?

The vertical, no less than the horizontal and seasonal
distribution of the Copepoda, must be of considerable
importance in connection with their value as fish food.
Some species, such as Temora longicornis, appear to live
only at or near the surface, while others, ike Hucheta
norvegica, are seldom found in less than 100 fathoms in
depth. Some species evidently have a vertical migration,
though what is the positive reason for this it would be
difficult to answer. Light appears, in some cases, to act

SURFACE AND DEEP TOW-NETTING. 157

as an attractive influence, in others as a repellant ; and it
is not improbable, as evidence seems to favour, that dif-
ferent intensities or kinds of light, whether as sunlight,
moonlight, or electric light, have various degrees of
attractive or repelling power.

At a depth of 33 fathoms, when about six miles west of
Peel, a cast of the steamer’s large trawl was made, a finely-
meshed tow-net being fixed to the line at about 3 fathoms
distance from the trawl. As the vessel slowly proceeded
the tow-net would be within a varying few feet from the
sea bottom. Simultaneously a similar tow-net was. let
out at the stern of the ship, and allowed to run at the
surface and to within a few feet thereof.

The temperature of the surface water was about 44°
Fahr., and only a slight ripple on the sea, and a cloudy
sky without sunshine.

Experience has often annoyingly shown us the great
risk to which a bottom tow-net, attached to a trawl or
dredge, is exposed from its passing over rough ground,
the tow-net frequently being either bodily carried away
or torn in such a way that the entire contents have
escaped. It was, therefore, with much satisfaction that
on this occasion when the trawl was hauled up after 17
hours immersion, the vessel steaming very slowly mean-
time, although the trawl-net itself was torn, the tow-net
was found to be in perfect condition, and containing a
considerable quantity of living plankton.

At the same moment the surface net was taken in, and
the contents of each net were immediately preserved in
bottles containing a 5 % solution of Formol and sea-water.

The following table is a summary of the respective
contents, Copepoda forming, as might be expected, by far
the largest proportion of each.

Allowing the contents time to sink the amount of

158 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

surface material was found to be 8 cc., that taken by the
bottom tow-net 20 cc., showing the latter to be 23 times
as much in quantity as the former.

TABLE showing Plankton taken by surface and bottom
(33 faths.) tow-nets, January 29th, 1899 :—

Surf. Townet | Bot. Townet

Calanus firmarchicus - - - F A
Pseudocalanus elongatus - - A A
Bradyidvws armatus- - - - | 5S
Centropages hamatus - - - C
& | Temora longicornis - - - - EF
a Metridia longa - - - - - F
&| Acartia claustt - - - - = A C
© | Candace pectinata - - - - | i
Oithona spinifrons - - - - C #
Euterpe acutifrons - - - = C
Longipedia coronata - - - | 5
Diatomacese - - - - - -/| A )
Foraminifera - - - - - - F
Meduse - - - - - - - F
Ctenophora. Plewrobrachia - FE
Chetognathe. Sagitta - - A
Crustacea, Amphipoda - - - EF
i larval Cirripedes’ - FE
Bs 5, schizopods - F C
Mollusea, larval - - - - - F
Bishsovas (ses) Seta F
Annelids, larval - - --- - FF F

A=abundant. C=common. F=few. S=scarce.

An analysis of the above table shows that four species
of Copepoda were common to surface and bottom, viz.,
Calanus finmarchicus, Pseudocalanus elongatus, Acartia
claus, and Oithona spinifrons. In proportion to the

SURFACE AND DEEP TOW-NETTING. 159.

amount of plankton in each tow-net, Calanus jfinmarchicus
was far more abundant at the bottom than at the surface,
the preponderance of males being very marked. Pseudo-
calanus elongatus was very abundant in both tow-nets, the
number of specimens in the lower one being equal to that
of all the other species put together.

Acartia claus and Oithona spinifrons were in about
equal proportions; more plentiful in the surface net than
in the bottom net, but few specimens of Ozthona being
found in the latter.

Four species of Copepoda were found in the surface
tow-net only, viz., Temora longicorms, Centropages hama-
tus, Longipedia coronata, and Huterpe acutifrons. The
two first-named are very common British species, being
rarely found at any distance below the surface.

Longupedia coronata is also common, though usually
found in rocky pools attached to fuci, between tidal limits.
The occurrence of Huterpe acutifrons in considerable ~
quantity is rather surprising, inasmuch as it 1s a rare
species round our coasts, and seldom found in any quantity.

It is of wide distribution, extending from South America,
into the Atlantic, the Canary Islands, throughout the
Mediterranean, the northern coasts of France, and at
Heligoland. ,

Three free-swimming species, not occurring in the sur-
face tow-net, were found in the bottom net, viz., Bradyi-
dius armatus, Metridia longa, and Candace pectinata, the
latter being entirely new to the L.M.B.C. district, and
the two former seldom met with.

Bradyidius armatus I have found on two or there
previous occasions at similar depths. It is readily dis-
tinguished from the common Pseudocalanus elongatus,
from which genus Giesbrecht has lately removed it, by the
strong spinous terminations to the cephalothorax.

160 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Metridia longa is decidedly rare in our district. Itisa
distinctly northern species, having been reported from the
Arctic Sea, Greenland, Spitzbergen, Scotland, and as far
_ south as the English Channel, and about the Channel

Islands. I have lately found it very plentiful in plankton

collected about Valencia, on the West of Ireland, and it
was frequently present in the collections taken by Prof.
Herdman in his traverse of the North Atlantic.

The finding of several fine specimens of Candace pec-
tinata, for the first time in our district, is notable. This
species was first found by Drs. Brady and Robertson, at a
depth of 40 fathoms, off the Scilly Islands. I have on
several occasions found it about the west coast of Scot-
land, and Mr. Thomas Scott reports it from the Frith of
Forth. I found it very sparingly off the Channel Islands,
and fairly plentiful about Valencia, Ireland. It thus
appears to be probably generally distributed round the
British Isles, and both at surface and sea bottom. Dr.
Brady reports the ‘‘ Challenger’’ captures of this species
about Australia, the Phillipine Islands, and between
Ascension and the Azores. I found it common about the
Canary Islands. It and the other members of the genus
Candace are at once distinguishable by the dark-coloured
antenne, spines, and plumes, and the terminal spines and
the swimming feet.

The surface Diatomacez were very numerous, while the
very few found in the lower net were probably taken
in as it neared the surface. Mr. Thomas Comber, F.L.S.,
to whom I sent the surface Diatoms for examination, has
kindly drawn up the following list of 18 species found,
V1Z. :—

DiaToMs—

Biddulphia mobiliensis, common.
Coscinodiscus-concinnus, common.

SURFACE AND DEEP TOW-NETTING. 161

_C. centralis, occasional.

Nitzschia paradoxa, frequent. .
Fragilaria hyalina, occasional.
Synedra Gazlloni, occasional.
Thalassiosira Frauenfeldii, occasional.
Hunotia sp., occasional, a fresh-water form.
Ehabdonema adriaticum.
Grammatophora marina.

Navicula gregaria.

Isthmia enervis.

Paralia sulcata.

Cyclotella striata, var. ambigua.
Chetoceros boreale.

C. pelagicum.

C. secundum.

C. fucellatum.

The few Foraminifera found were at or near the surface,
as also were the few Meduse taken.

Several Pleurobrachia were in the bottom net, but none
in the surface; and an enormous number of Sagitta were
brought from the bottom, but none were found near the
surface.

A few fish ova were taken by the bottom net only. The
distribution of the remaining plankton taken can be seen
from the table (p. 158).

From these comparative results of the gatherings taken
from upper and lower strata, it is clear that both should
be noted at any given time, as there is much to be yet
learned as to the effects of temperature and other influences
upon the minute forms of life of our seas. It is hoped
that we may be able, say at the four seasons, during the
rest of the year, to continue observations on a similar plan.

To obviate the difficulty of arranging a bottom tow-net,
there is the alternative method of pumping water directly

a '

162 TRANSACTIONS, LIVERPOOL BIOLOGICAL SOCIETY.

up, from any depth, through a hose pipe sunk as low as ©
required, and worked by a hand pump fixed on the —
steamer’s deck, the water then passing through an ordinary _
fine tow-net. This has been already successfully tried by —
Dr. Kofoid and others in America, and recently by Mr. 4
Garstang, from Plymouth. A suitable pump and hose ©
has been supplied to the Port Erin Biological Station —
for use on the next and all future expeditions, ween the —
results will be duly recorded. | P

163

SOME SPECULATIONS on the DERIVATION of
our BRITISH COLEOPTERA.

By W. EH. S#Harp.
[Read January 13th, 1899.]

Amone the many problems to which the attention of the
modern Biologist is invited, that of the present distri-
bution and probable derivation of floras and faunas is one
of the most interesting and fascinating—interesting because
in the most cursory survey of the subject so many issues
are seen at once to be involved; fascinating because so
wide a door is thrown open to the most unfettered
theorizing, and because in any adequate comprehension
of these problems possible answers to so many questions
seem to lie hidden—questions propounded by the Geologist
not less than by the Biologist.

No one will dispute that this enquiry is a very difficult
and perplexing one; that its factors are more than usually
complicated and obscure; and that the evidence we have
available is, as it affects vast groups of animals, meagre |
and defective. As Dr. Wallace has well said in a recent
work :—

“There is no short and easy method of dealing with
these questions (that is, the present distribution of organic
life), because they are in their very nature the visible
outcome and residual product of the whole past history of
the earth. If we take the organic productions of a small
island or of any very limited tract of country, such as a
moderate sized parish, we have in their relations and
affinities—in the fact that they are there and others are
not there—a problem which involves all the migrations of

164 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

these species and their ancestral forms—all the vicissi-
tudes of climate and all the changes of sea and land which
may have affected these migrations—the whole series of
actions and reactions which have determined the preserva-
tion of some forms and the extinction of others—in fact,
the whole history of the earth, inorganic and organic,
throughout a large portion of geological time.”

Now to invite your attention to so vast a subject as
this will, I am sure, appear a considerable presumption
on my part, and I will hasten to alleviate your apprehen-
sions by saying that my intention is only to consider one
order of one class of the British Fauna, and I do this
with the more confidence from the fact that of the distri-
bution of the British Coleoptera very little is really
known, and of its proximate derivation still less. It
therefore appears a safe topic on which to theorize, our
ignorance of facts allowing us a more unbiased freedom in
speculation. Apart, however, from this rather dubious
qualification, to render such an enquiry as this—that, is,
the distribution and origin of the British Fauna—manage-
able, it becomes necessary to narrow the question down
to one small group of that fauna: one must attack it in
detail if one wishes to do so at all exhaustively. Further,
it appears to me that the Coleoptera-or Beetles afford
particularly good material for such an enquiry, and here I
am not indeed sure whether it is not perhaps necessary to
defend the cause of the terrestrial invertebrate in limine,
so strong are the traditions of this room in favour of the
marine—so permeated is its very atmosphere with ‘‘ mur-
murs and scents of the infinite sea.”’ Butit seems obvious
that when we come to talk about distribution your student
of terrestrial life has most to say. Fresh-water faunas are
another matter, but I venture to think that not much as
regards past earth movements, submersions and upheayals,.

:

DERIVATION OF BRITISH COLEOPTERA. 165

streams of migration and so on, can be deduced from a
study of purely pelagic life.

Thrown back, then, on our terrestrial organisms, and
leaving out of account the flora which, in many ways,
affords better and more abundant evidence than does any
part of our fauna, we find the mammals and reptilia far too
small in specific numbers to be of much service in the
enquiry. There are certainly in the distribution of our fresh-
water fishes most valuable and significant facts; but it is
to the invertebrates which far out-number all the other
organisms put together that we find our most abundant
material and safest guides; and of the invertebrates,
certainly the insects have been most closely and generally
studied, and of them consequently the British distribution
is perhaps best known.

The majority of insects however fly, and, of course,
the evidence afforded by creatures whose transport is so
easy may readily become misleading; but of all the insects
the Beetles use their wings least; in fact, some of them
do not possess these organs in any effective degree. The
Coleoptera too, are exceedingly numerous, both specifically
and individually, some 3,300 species being recorded as
British, and they are perhaps more uniformly and generally
distributed than any other order of insects.

I will therefore invite your attention to what is known,
and still more, to what is unknown, of the distribution of
the Coleoptera of these islands and their possible derivation,
premising only that any inferences we may draw from
such facts, unless equally applicable to the distribution of
all the other fauna and all the flora as well, are theoreti-
cally invalid.

Now the first broad fact to bear in mind is that the
whole British Coleopterous Fauna, and, indeed, we might
say the whole British Fauna, is simply part and parcel of

166 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

that of north-western Europe. It is merely a restricted
extension of what is known as the Palearctic Fauna,
possessing, at any rate among the Coleoptera, hardly any
distinct forms of specific and very few of even varietal
value. I am quite aware in saying this that there are
some 67 species out of our British total which so far have
not been publicly recorded from any other country. I
venture, however, to dismiss these as evidences. of any
distinct Coleopterous Fauna, because, firstly, the bulk of
them are merely of slight varietal value; secondly, they
are all exceedingly rare, many resting as unique specimens;
and thirdly, because I can recognise no presumption that
because they have never been recorded, they therefore
necessarily do not occur elsewhere in Europe.

If, then, we seek the phylogenetic origin of our British
Beetles, we must find it in the origin of the whole Coleop-
terous Fauna of the north temperate zone, and here we
become at once lost in the dark abyss of Geologic time.
We certainly know that insects, as we find them now, are
of vast antiquity. As far back at least as the carboniferous
forms referrable to existing orders appear, while fossil
Coleoptera generically allied to present forms have been
found in Oolitic and Cretaceous deposits in England.
Indeed, it seems safe to assume that the bulk of our
present species of insects peopled the England of, at any
rate, later Tertiary times. The evidence of this is sup-
plied by Beetle remains, wing cases, and so on, found in
the Cromer Forest beds, which can be almost specifically
assigned to modern Harpali and Donacia. These beds,
I believe, are generally admitted to be, at any rate, ante-
glacial, and the remains in them probably represent the
late Pleistocene fauna-and flora of the British Isles, if you
can so term some indefinite extension of north-western
Europe, of whose configuration.and limits -we can. only

:

__ DERIVATION OF BRITISH COLEOPTERA. 167

vaguely guess. Further back than this we cannot go, the
Geological record as regards insects is extremely fragmen-
tary and discontinuous, and we really have not the slightest
evidence as to when, where, or how the differentiation of
existing specific forms among the Coleoptera, or indeed of
any other order of insects, took place.

But we are not now considering the origin of insects,
but their distribution in Britain and immediate derivation,
and we cannot get far into the subject before we discover
that any theories we may form on the matter will depend
ultimately on the view we take of that vast secular catas-
trophe known as the Glacial Age.

How intense was that cold? Did an ice sheet cover the
whole of the British Islands as to-day it does Greenland ?
Can we regard the almost entire submergence of this
country during or at the close of the glacial epoch as
incontrovertible? In short, were the climatic and other

‘conditions such that the whole of the preceding Tertiary

fauna of Great Britain must have been extirpated and
destroyed? Was such a clean sweep made of the whole of
antecedent life that when the great refrigeration passed, or
when the land stood once more above the waters, there
was presented a tabula rasa, and an empty stage on
which the actors in our present drama of life might appear
for the first time ?

These are the questions which, at the outset of our
enquiry, confront us, questions you will observe so very
much more easily asked than answered. For it appears
that opinion on these points is not quite uniform, and,
indeed, seems recently to have become, as regards the
severity of this refrigeration, somewhat modified.

Many great authorities, however, hold extreme views.
Thus, Dr. Wallace says—‘‘The submergence destroyed

168 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,

this (ante-glacial fauna) or at least the greater part
OL ie

Prof. J. Geikie remarks—“ As neither our animals nor
our plants could have existed in the British Islands during
the last glacial epoch, it follows that they must be of post-
glacial age;+ and another Geologist, Mr. Clement Reed,
says—‘‘In the Britain of the present day we may study
the repeopling of a country over which everything had
been exterminated.’’t

On the other hand, Dr. Scharff, of Dublin, has lately
written a very interesting monograph on this subject,
the burden of which is to demonstrate the probability of
the uninterrupted continuity of the British Fauna since
pre-glacial times. For so revolutionary a view, however,
the evidence seems inadequate. The climatic conditions
which we know from data quite other than faunistic must
have prevailed, still more such a submergence as the
shell-beds of Tryfan and Macclesfield might suggest, could
not but have very considerably modified the previous
fauna and flora, if it did not exterminate them. A
remnant may have escaped the cold by a southern migra-
tion, and an archipelago of islets may have afforded a
refuge from a submergence; but I doubt whether, in any
case, 1t can be safely maintained that our present flora
and fauna are merely an uninterrupted continuation of
those of late Tertiary days.

However, leaving ground so debatable, let us turn to
the animals themselves, and see what light they throw on
the subject.

In the first place, we must exclude from consideration
any species which owes its appearance here to other than
purely natural agencies. There are probably very ‘few

* “Island Life,” p. 338. t ‘*Gr. Ice Age,” 3rd ed., 1894.
t “ Flora of Greenland.”

'

DERIVATION OF BRITISH COLEOPTERA. 169

such introductions in the British Fauna, and more in the
flora than the fauna of most islands. By non-natural, I
mean introductions which are in any way directly or
indirectly due to the appearance of humanity on the
scene. Cases of the sort, of course, do occur, and impress
one by the rapidity of their advance. There is, for
instance, the oak-gall fly, Cynips kollart, which produces
our familiar oak apple, and this insect appears to have
been first introduced into Devonshire and restricted to
that county not much more than half a century ago—now
every oak tree in the country testifies to the extent of its
present range. The deplorable ubiquity of the common
cockroach is another case in point among the insecta.

On the other hand, among mammals we have the poor

success which has attended all attempts to introduce our

common English hare into Ireland, whose native form is
a closely allied but distinct species the Arctic or moun-
tain hare. Here some unknown influence seems to resist
the establishment of a new species in an area which appears
to be perfectly suitable to its maintainence. Generally
speaking the adjustment between existing species and
their environment in any region is so close and so exact
that there is no room, not the least chink of an interstice,

into which a new arrival can thrust itself, and the nearer |

genetically such a would-be immigrant may be to forms
already existing, so much the more difficult is its entrance
made.

Where we do find such introductions 1s where the
environment itself has been changed by man, and among
the Coleoptera there are perhaps 40 or 50 species, quasi-
domesticated, the Beetles of granaries and bakeries and
grocers shops. These we must eliminate in any considera-
tion of the natural distribution of the group.

But, considering the matter a little further, we discover,

170 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

from a very brief study of the physiological characteristics
of the Coleoptera, as of any other order of insects, that the
species themselves clearly fall into two groups, which we
may call the adaptables and the non-adaptables, or, if we
like, the progressives and the conservatives— that is,
species which are common everywhere, and species which
are either generally rare or common only locally ; in other
words, there are a certain number of forms which have
successfully solved the problem of the continual adjust-
ment of themselves to an ever-changing environment, and
those who, failing to do so, perish when their own par-
ticular environment ceases to be. These latter certainly
form the bulk of the species. There are Beetles attached
to the sandy wastes of the shore, to marshes, forests,
heaths, mosses; and when you turn your sandhills into
docks and golf links, drain your mosses and bogs, cut
down your woodlands and cultivate your desert wilder-
nesses, then the non-adaptable, the conservatives of the
feral population of such places, pass to return no more;
they are incapable of the necessary new adaptations.
Hence it is that inasmuch as the physical characteristics
of this country are continually changing, and that at an
increasing ratio, the extinction of most of our conservative
species of animals is merely a matter of time. This pro-
cess has almost completed itself among the mammals, is
in full swing among the birds, and is quite noticeable
among such an insect group as the Diurnal Lepidoptera.
It must, however, be admitted that there is no evidence of
any extinction of specific forms among the Coleoptera as
yet, and I have, perhaps, rather exceeded the strict lines of
“my subject on this point, because I want to impress the
distinction between such species, which are the only ones
that are of any service to us, in our endeavour to discover
the derivation of any of them—between these and those

DERIVATION OF BRITISH COLEOPTERA. 171

other progressive species who have established a sort of
modus vivendi with civilization, readily adapt themselves
to those altered conditions which agriculture, among other
agencies, has imposed, and thrive under a new regime.
Such species we call common, and most of the others rare,
but the mystery of why one species of a genus can so
adapt itself, and another, so far as we can see, similar
physiologically, refuses to do so, is as impenetrable as
ever; and to say that the fittest always survive only
restates the fact in other words, and does not explain it.

However, the point is that those species which have
become fit, and are therefore common everywhere, are not
the slightest guide to us in our search for the causes and
methods of original distribution. Their facility for adap-
tation is so great, that whether they may have arrived
early or late, from the north, south, east, or west, it comes
to quite the same thing, that is, their present ubiquity
and the complete obliteration of all their past proceedings.

We must thus eliminate at the commencement of our
enquiry all such quasi-domesticated Beetles as those of
the genera Dermestes, Pellio, Anthrenus, or Alphitobius,
and all such ubiquitous species as Nebria brevicollis, or
Harpalus eneus, which appear to be equally abundant
from Cornwall to Caithness, and from the top of Ben
Nevis to below high-water mark.

These exceptions, however, will be found to be but
trifling, and when we study the distribution of the great
bulk of our Coleopterous Fauna which will remain, we
shall, I think, discover that they can be, with more or less
precision, arranged in at least three groups, or rather, that
in their grouping we discover at least three diverse elements,
and this diversion will.be quite independent of genetic
affinities ; in fact, we shall be able to find in any of the
larger genera members belonging to each of them.

172 TRANSACTIONS LIVERPOOL BIOLOGIGAL SOCIETY.

We have, firstly, a somewhat small group of species
occurring in regions of the north and north-west, elsewhere
inhabiting only mountains and elevated moorlands. This
group has been called the Arctic, the Glacial, or the Celtic
Fauna, all terms which, to some extent, beg the question
at issue. Perhaps the Celtic is the better term, if we can
eliminate the idea that the group is necessarily comparable
with the Celtic race, ethnologically considered. Its range
is principally in Scotland and western Ireland. Several
species occur there which do not occur in England, .but no
single species occurs in England which does not also occur
in Scotland or Ireland, or both. Next we have a group
which contains by far the greater proportion of our species.
They have been called the Szberian immigration ; whether
their ultimate starting point was Siberia or not, it is clear
that their proximate origin was the plain of Central
Europe, that they arrived here from the south-east and
east, and that during their dispersal throughout this
country some obstacle, though probably not entire separa-
tion, prevented complete access into Ireland. These are
species generally but by no means equally distributed over
Great Britain, not restricted to elevation, but often rigidly ~
so by other conditions of environment; from a maximum
density in the south and east, they thin out northward
and westward, till many of their forms are rare in Scotland
and some altogether wanting in Ireland.

The third group is perhaps more difficult to define, and
its members cannot always be disentangled from those of
the second group. We should comprise in it all those
species whose distribution is exclusively southern or south-
western, and is often marked by extreme discontinuity.
his is known as the Teutonic element in its eastern and
the Iberian in its western extension. The former, the
south coast of England, the latter, the south and south-

DERIVATION OF BRITISH COLEOPTERA. 173

west of Iveland. The characteristics of this fauna are
very enigmatical; some species occur only in England,
some few only in Ireland, and some are common to both.
From Scotland the group is quite absent.

Now, from what is now known of the distribution of the
British Coleoptera, we could go through the entire list
and allocate, with more or less precision, nearly every
species to one or other of these groups. And the questions
suggested by so doing would be something lke these—
Do such groups really represent separate waves or streams
of immigration? Are they of different age, and if so, in
what relation as regards time do they stand to one another?

_ Did separate areas originate them, or only separate

epochs?

Now, to answer, as some have done, firstly, that all life
was destroyed over these islands either by the severity of
elacial conditions or by submergence, and secondly, that
during the slow subsequent amelioration of the climate
these streams of migration arrived in the order in which
I have mentioned them, and that sometime during this
process, but subsequent to the dispersal of the first group,
the connection between Great Britain and Ireland broke
down, may appear a simple explanation, but however well
it may fit some of the fauna or flora, certainly it does
not exhaustively explain all the facts of their distribu-
tion. It does not explain the vestiges of a distinctively
Iberian fauna, still less those of a North American one in
Ireland. —

To make this clearer, let us consider these groups a
little more in detail. Our first or Celtic division’s maxi-
mum development is in the north, from Orkney and
Shetland, over all the Highlands and all the western isles

of Scotland, and all the mountains north and west of Ireland

174 TRANSACTIONS LIVERPOOL BIOLOGIGAL SOCITY.

down to Kerry. It is only partially represented in the
Welsh mountains, and some of its species occur spora-
dically in England. In Europe all the species occur in
Scandinavia, many of them on the Alps. Take two of
them—that large water Beetle, Dytiscus lapponicus, and a
smaller ground Beetle, Pelophila borealis. Both occur in
the extreme west of Scotland and Ireland, where they are
abundant, but neither in England, although in Cumber-
land, Yorkshire, or Wales the environment would be
perfectly appropriate to them. Now, it is difficult to
understand, if these two species formed part of the earliest
immigration from the south-east or east, why they should
be both absent from an area over which they must have
travelled to arrive at their present home. _

Other species, such as Carabus glabratus and Miscodera
arctica do occur at suitable elevations in the north of
England and in Wales, while one species, Carabus clath-
ratus, with a general range almost as limited as Dytiscus
lapponicus, has been recorded from Suffolk, but there is
no example of a high mountain or moorland species which
attains a maximum in England or Wales, and thence thins
out north or west, such as one might have expected had
the original entrance of such species been from the south
or east at however remote a date.

Leaving the Coleoptera however for a moment, but still
considering the Irish fauna, we cannot but be struck by
the remarkable occurrence of three species of North
American fresh-water sponges in the south and west of
Ireland, species not occurring elsewhere in Europe. These
were discovered by Dr. Hanitsch—a well-known former
member of this Society—about four years ago.

Dr. Hanitsch himself suggests that their presence might
be due to the gemmules of these sponges having been
carried attached to the feet of birds across the Atlantic.

a ee Oe

DERIVATION OF BRITISH COLEOPTERA. 175

They may also afford evidence of a former land connection
which, closing in the Atlantic on the North, united
Treland, Scotland, Norway, Iceland, and Greenland. Such
a continental land extension might well have been the

home of what we call the Celtic element in our western

fauna, as well as the bridge which carried those American
species which we recognize both in the Irish flora as well
as fauna. Thence any immigration would have been from
the north-west, south and eastwards, which is what the
facts of our Coleopterous distribution of this group require.
The difficulty comes in when we attempt to harmonize
this theory with preconceived notions of the severity of
glacial times, for, of course, if that severity were such as
to forbid the existence of animal life in Great Britain,
still more would it be impossible in this hypothetical
northern land, and any migration would necessarily have
had to proceed from the south northwards. If, however,
we may be allowed to suppose that the British glacial
chmate after all might have permitted the continued
existence of this Celtic fauna, then we see no difficulty
in conceiving that fauna, as originally a pre-glacial one
inhabiting a region to the north-west, now lost and driven
southward by the advent of glacial conditions which cer-
tainly would have made such a northern land untenable,
instead of led up northwards from the south on the
cessation of such conditions. In any case, I think we
may look upon the species comprised in this first group

as distinctly older residents than the vast mass of species

whose advent was more indubitably from the south and

east, and perhaps this is as much as one can safely say.

Proceeding, then, to a consideration of our second
group, we see no reason to doubt that their advent was
from the east and south-east at a time when the German
Ocean was dry land. The distribution of some of them is

176 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

still very limited ; thus, I should assign to this group such
an insect as Nebria livida, whose occurrence is restricted
to the sea coast of Yorkshire, and most of the fen species.
Many of them are phytophagous and attached to special
plants; others, creophagous or necrophagous, and so
dependent on the existence of higher animals. These
factors rather complicate the question, as it becomes
evident that the distribution of the Beetles is limited by
the possibilities of distribution of their various hosts and
food plants. I have already remarked that a proportion
of this group does not seem to have reached Ireland at
all. It is quite true that up to within about ten years ago
very little was known of the Coleoptera of Ireland, and
that, since that time, every year has added species to the
Irish list, but as the Irish deficiency in mammals, and
still more in reptilia, is certain and notorious, it is
perhaps a fair presumption that many of these Coleop-
tera do not really exist there. In this connection I should
like to point out that the Manx Coleopterous fauna, so far
as it has been studied, is quite Irish in character; even
that peculiarly Irish form Szlpha subrotundata occurs
there. It exhibits the same preponderance of the Celtic
group, the same differences in the eastern one, and,
curiously enough, a quite southern English species, the
well-known rose chafer, Cetonza aurata, which has rather
surprised entomologists by having been recently found in
Arran Island, off Galway and in the extreme west of
Kerry, Clare, and Donegal; has also occurred in the
Isle of Man. So little, however, is really known of the
Manx Coleoptera, that 1t would be rash to hazard any
theories on the meagre facts in our possession.

We may, I think, take it for granted that no species
now, except under exceptional and non-natural conditions,
is Increasing or ever will increase its range. No doubt

DERIVATION OF BRITISH COLEOPTERA. 177

freedom from competition, either directly or indirectly
through food plants, &c.,in certain suitable areas first
fixed the lines of migration and settlement. As however,
the available land surface became more and more crowded,
fixity of tenure would become more and more the order of
things, so that it seems probable that there must have
been a point somewhere in the past which marked the
final equilibrium of the forces of dispersal and the stress of
competition, and that since this point was reached, most

- movement has been in the form of retrogression and due

to human interference with environment.

We can thus understand that, given an original exotic
immigration of an assemblage of species from the east
(firstly), the rate of progress, initially perhaps high, would
be continually decreasing till it came to a dead stop, and
that no length of time elapsing since then would renew it
except by organic change in the species themselves through
natural selection, and that (secondly) the rate of advance
among the species themselves would be very unequal. In
Nature the race is always to the swift, the battle invariably
to the strong. This explains why some species arrived in
Ireland while others were too late, and so cut off, or, what
is more probable, since there are similar deficiencies in
the Scotch Fauna, that their rate of progress was so slow,
that before they had got half this distance, progress at
all became impossible.

The characteristics of this second or Central Huropean
group, then, are that from a maximum in the east and
south-east it thins out both specifically and individually
westward and northward, and if we find a species, as we
do several, whose maximum seems to be in Kerry, Corn-
wall, and South Wales, but is absent or rare in Norfolk,
Essex, and Kent, then we may, I think, assign such a
species ipso facto to our third group.

178 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

- To a consideration of this assemblage we will therefore
now proceed, and it undoubtedly exhibits perhaps the
most puzzling features of them all. I should include in
this group, firstly, all species whose northward range does
not pass beyond the Thames Valley and the Bristol
Channel, and secondly, a certain number of Southern
Kuropean forms found in Ireland, but not to the same
extent in England. There can be no doubt but that this
element in our fauna is distinctly southern, or relative to
Hurope as a whole south-western. There seems to have
been two lines of migration, one, which crossing the
English Channel from directly south, does not appear to
have extended further westward than Devonshire, another
which seems to have had an origin in some much more
western ancient land extension, spread itself through
Ireland in a curiously disconnected manner, and leit
vestiges of its migration in Cornwall and Devonshire.

Calling this then the southern group, we find :— |

Firstly, species such as Lucanus Cervus, Cicindela ger-
manica, or Carabus intricatus among the Beetles. Among
other insects, the Giant Earwig, only known to inhabit a
few isolated spots along the Sussex and Hampshire coast,
or the butterfly called the Lulworth Skipper, found only at
Lulworth, in Dorset. These are species which do not
extend northward of the Thames Valley, are very rare or
absent west of Dorset, and quite absent from any part of
Treland. |

Secondly, species common to the south and west of
Ireland and England, but principally the extreme west of
England; such are Mesites Tardyi and Chrysomela Banksit.

Some of these exhibit discontinuity of range to so high a
degree as almost to suggest two separate streams of migra-.
tion. Thus Pyropterus affinis, a brillant scarlet Beetle and
one not likely to be overlooked by collectors, has so far been

DERIVATION OF BRITISH COLEOPTERA. 179

taken only in Sherwood Forest and Killarney. That bril-
hant insect, too, Lytta vesicatoria, until, to the amazement
of coleopterists, 1t turned up last year in Roscommon, might
have been regarded as a peculiarly English species, as it
had only been taken a few times in Kent and Hampshire.
Another conspicuous Beetle, Nebria complanata, in Eng-
land, inhabits only the sandy shores of the Bristol Channel,
and, in Ireland, a short strip of the Wexford coast
immediately opposite. This case however, I consider it
allowable to hold to be due to accidental transference,
since the species is purely littoral in its habits and is fond
of hiding under and in the crevices of drift wood lying on
the shore. Hence its transport across from the Bristol
Channel to Wexford is not inconceivable.

And thirdly, we have species which are only found in
Ireland; this has been called the Iberian element in the
fauna of that enigmatical island and the spotted slug of
Kerry. Geomalacus maculosus is a famous example of it.
It is, I believe, also well marked in the flora. There is a
house spider, Tegenaria hibernica, the special pride of
Dublin, and a jumping spider, Attws floricola, from Lough
Corrib, discovered only two years ago, both unknown else-
where in the British Islands. Among the Coleoptera, so
far, we can name only Stlpha suwbrotundata and Otiorr-
hynchus auropunctatus, and, curiously enough, this latter
species has more an eastern than a western range; but
when the wilds of Kerry and Clare have been more
thoroughly explored by Coleopterists, I doubt not that the
list will be larger one.

Now, these facts require a great deal of explanation, the
more so if we grant any community of origin to the species
under discussion, for some of them exhibit great discon-
tinuity of distribution which, according to Dr. Wallace
and other high authorities, infers great antiquity, of origin,

-180 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and others an equal but limited homogeneity, which seems
to imply a recent arrival.

I think we must admit from their generally southern
range that they are all post-glacial, and it might also be a
plausible conjecture that the Iberian element, that is, the
Trish or Irish and West Anglian species were earlier than
the South Anglian. It seems also probable that while it
is certain Ireland must have been joined with an extreme
western extension of Europe during the advent of the
former, that country may have been quite insulated while
the advent of the latter took place across what 1s now the
eastern part of the English Channel. Moreover, there
seem to be grounds for assuming that at a time when
Cornwall, Kerry, France, and Portugal were all united,
some obstacle, such as an immense lake, existed where
the Irish Sea and St. George’s Channel are now. ‘This
would explain the phenomenon of bifurcation in the lines
of distribution of such a species as Cetonta aurata, which
does not extend much beyond the midlands in England,
while it goes right up to Donegal in Ireland, and also
occurs in Man; or of Pyropterus affinis, which, as I
have mentioned before, occurs in Kerry in Ireland and
Nottinghamshire in England.

Further than such problematical conjectures, or, at
best, working hypotheses, I doubt if we are justified in
going for the subject is a very difficult one, and in
my own opinion hardly at present ripe for discussion.
We have after all not a very complete basis of facts to go
upon, there may have been many streams, and many
periods of migration from the same region, and I doubt
whether it is possible, perhaps ever will be possible, to
quite disentangle these threads, or to prove the exact
limit as regards species of northern, eastern, or southern
derivation.

DERIVATION OF BRITISH COLEOPTERA. 181

Indeed, none of these various schemes, conjectures, and
hypotheses will stand working out in complete detail. All
we can hope to do, at any rate at present, is to get hold
of some few general principals, and to test those by all the

evidence at our command, and by application to the whole

of both fauna and flora.

Some such principles I hope emerge from the fog of
conjecture through which we have been travelling. As
that we can certainly detect a threefold, if not a fourfold,
disparity in our insect fauna; that its phenomena point,
at any rate, to past land extensions to the north-west
and to the south-west, now entirely swept away; that
Ireland must have been separated from the greater part
of England by an obstacle of some kind while the fauna
was in process of establishment, and while or after it was
itself continuous with an extension of Western Europe,
and that it is not impossible that some extreme estimate
of the severity of the glacial climate may require modifi-
tion. Furthermore, that if any of our Coleoptera can be
termed autochthonous, the northern or Celtic element is
the only one that can be so regarded, and that, while all
the rest of the fauna seems to have been derived from the
south and east, this only may have had a northern origin
and be possibly pre-glacial. ‘These are conclusions which
we may perhaps entertain as provisional hypothesis. For
the rest, perhaps all I have succeeded in making plain is
how vastly our ignorance exceeds our knowledge of the
factors of the problem; and this is especially the case
with a group of insects such as we have been considering.
For mark the difficulties of the quest. Insects, although,
because they are probably much older as distinct species,
and also because both specifically and individually they
are very much more numerous than the higher vertebrata,
furnish more abundant evidence of faunistic distribution,

182 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

yet because of their minute size and curiously spasmodic
appearances and disappearances make the investigation
of their occurrences, and more especially of their non-
occurrences, all the more difficult.

For instance, whether or no the beaver is now extinct
in the British Islands, there can hardly be too opinions.
Scarcely the most persistent optimist can cherish the
hope that the great copper butterfly will ever again be
seen of mortal eyes in the eastern fens; but what amount
of negative evidence is sufficient to justify the assumption
that such and such a species of Homalota or Atonaria
does not occur in any particular locality. A single pro-
perly verified record is sufficient to establish the positive
side, but as the proof of the negative is proverbially a
hard matter, so in a case where the subjects are so
minute and so elusive, it verges on the impossible.

Furthermore, there is an even still more perplexing
factor in the problem, and that is what I have just alluded
to as the spasmodic method of appearance of so many
insects. This is a very interesting subject, and very
worthy of more attention and investigation than it has
yet received. The extraordinary irregularity of such an
insect as the hawk moth, Deilephila galw, is a case in
point, but the characteristic prevails more or less among
all insect life. | |

Doubtless the meaning of such phenomena is to be
found in the singular delicacy of adjustment between
insects and their environment, so that the most minute
and obscure change in the latter is responded to in a
greatly enhanced and apparently exaggerated degree by
the former, although the connection may be quite unknown
to us, and often indirect.

But I firmly believe that a large proportion of what is

DERIVATION OF BRITISH COLEOPTERA. — 183

generally regarded as discontinuity of occurrence should be
more correctly attributed to discontinuity of appearance.

Consider, for instance, such a case as Aegialia rufa, a
small Lamellicorn Beetle supposed to be peculiar to the
littoral of Lancashire and Cheshire in this country. Now,
within say the last thirty years, this district, that is, the
coast sandhills from Southport to Crosby and from
Wallasey to Hoylake has been, perhaps, as well searched
year by year by entomologists as any other of equal extent
in the kingdom, and this perplexing insect has during
three or four summers of this period appeared in profusion.
On those occasions it has, of course, been discovered and
recorded for this district only. This, on the face of it,
supples an amazing instance of discontinuity of distri-
bution; but if the sandhills all round the coast had been as
persistently searched as those of Lancashire and Cheshire,
is it unimaginable that Aegialia rufa might, in its years
of plenty, have been taken in many other localities; in
fact, wherever round these coasts a suitable habitat offered
itself? You observe that, in this case, the Coleopterist
might have searched the Crosby and Wallasey sandhills
seven or eight years in succession, and declared as the
result of his investigation that Aegiaha rufa did not occur
in that district. Another observer goes there the next
season and finds the insect one of the most abundant of
the fauna.

You will gather from this that the study of the distribu-
tion of our insect fauna is beset with peculiar difficulties,
and that only close and systematic observation, extending
over a long course of years, is sufficient to justify the
assumption of the non-occurrence of many species in any
particular locality. This is just what we have not got. I
doubt if there be fifty Coleopterists in the whole of the
British Islands who systematically record the occurrences

184 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. .

of the order which they study. It is true we have a
sufficient mass of evidence to enable us to form broad
generalities, such as I have indicated in this paper; but
until that evidence is more copious and more complete, it
is obvious that they cannot be much more than pro-
visional, and it is certain that at present the aim of all
students of this—and, indeed, all orders of insects—should
be more the accumulation of facts than the exploitation
of theories.

I will therefore conclude these somewhat unsatisfactory
and disjointed observations by expressing the belief that
among the various functions of the many local natural
science and biological societies now in existence among
us, none excel in importance the accurate and systematic
verification and record of the flora and fauna of their own
particular district. As I have already said, many of the
feral inhabitants of these islands are swiftly passing; many
to a future generation of investigators will be but empty
names, and as they disappear, leaving no trace of their
sojourn here, they reduce little by little the possible
evidence by which we may ever hope to understand the
problems of their past migrations and distribution. Much
evidence of this kind is undoubtedly beyond recall, but it
is obvious to aiy one who has at all studied the distribu-
tion of our Insecta, that had we, instead of the disconnected
and often overlapping lists of records of individual students
and collectors, the authoritative table of corporate bodies
of workers, formed on some comprehensive system, care-
fully verified, and recorded in some permanent and pro-
eressive method, we should be enabled far more hopefully
to face the task of the elucidation of such a tangled web
as has formed the subject of our attention this evening.

Ce a ae ee a a

185

On SECONDARY THICKENING in the AERIAL
ROOTS of HEDERA HELIX.

BY
R. J. Harvey Gisson, M.A., F.L.S.
Professor of Botany in University College, Liverpool.

With Plate III.
[Read January 18th, 1899.]

I RECEIVED lately, through the kindness of Miss May
Rathbone of Neston, Cheshire, a portion of an ivy plant
whose aerial roots appeared to have undergone a some-
what curious change. Instead of showing the usual small
aerial roots of from half-an-inch to an inch in length, the
branch was provided with roots 6 or 8 inches long, pro-
fusely branched and considerably swollen especially for
the last 2 inches of their length.

The plant had been grown in a large pot in a room and
trained up the side of the window, and from about a foot
above the pot up to the topmost branches which twined
round the curtain rod exhibited both normal and abnormal
roots. They developed from the stems at all angles, quite
irrespective of light, and often all round the stem.

The normal aerial root of ivy shows, as has been
described and figured by Van Tieghem (L’origine des
membres endogénes, 1889) and others, a central vascular
cylinder surrounded by a well marked endodermis (d)
fig. 1. There are, as a rule, five protoxylems (/) united
centrally by a strongly sclerotic medullary region (9).
The pericycle (e) is, as a rule, from one to two layers thick
between the phloem and the endodermis, but may be three
layered opposite the protoxylems.

186 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The phloem (/) is small celled and not markedly distinct
from the pericycle.

The endodermis is followed by a fairly thick cortgx (c)
limited by a piliferous layer (a) replaced later by an
exodermis (0b). .

On examining the swollen roots one finds the number of
protoxylems greatly increased. A section of such a root as
that shown at figs. 2 or 3 may show from 12 to 20 pro-
toxylems united by a sclerotic medulla.

The protoxylems stand out prominently, creating deep
bays in which the phloem lies (fig. 4).

The pericycle is, in the swollen roots, much thicker,
averaging 4—5 cells in depth.

The endodermis is again well marked and is followed
by a thick cortex.

As the root becomes older two cambiums make their
appearance in a manner quite similar to that seen in a
normal terrestrial dicotyledonous root.

The first beginning of the change is apparently the
development of a phellogen from the most external layer
of the pericycle. ‘Tangential division in this layer results
in the formation of a distinct layer of phellem and is
followed by the shrivelling of the original cortex and
rupture and disintegration of it as well as of the endo-
dermis. At the same time, or more commonly somewhat
later, a cambium appears inside the phloem at the base of
each concavity formed by the prominent protoxylems,
becoming gradually completed from the pericycle out-
side the protoxylems. Secondary thickening, chiefly of
xylem, is now added in the ordinary way until the bays
become filled up with secondary xylem (figs. 5 and 6).

In the flattened roots (fig. 7) as many as 50 protoxylems
or even more (fig. 8) may occur.

AERIAL ROOTS OF HEDERA HELIX. 187

Similar instances of secondary thickening in aerial roots
are known to occur in certain Melastomaces, but I am
not _aware that the phenomenon has been previously
observed in Hedera; at all events I have been unable to
find any reference to the subject in the literature.

EXPLANATION OF PLATE.

Fig. 1. Transverse section of normal aerial root of
Hedera helix.

Figs. 2, 3and 7. Abnormal roots of Hedera.

Fig. 4. Transverse section of abnormal root, showing a
large number of prominent protoxylems.

Fig. 5 and 6. Development of phellogen and cambium
in abnormal roots. |

Fig. 8. Transverse section of abnormal root with 50
protoxylems.

-188

ADDENDUM to REPORT on a small COLLECTION
of ANTARCTIC PLANKTON,* &c.

SincE the publication of the paper with above title in last
year’s L. B. 8. Trans., I have received a communication
from Mr. William §. Bruce, F.R.S.G.S., who it appears
was along with Dr. Charles W. Donald, the collector of the
Plankton handed to me by Prof. D’Arcy Thompson, C.B.
Mr. Bruce points out to me that it was due almost entirely
to the generosity of Mr. B. Leigh Smith (of Franz Josef
Land fame), who contributed nearly £100. towards the
Biological outfit, that the collection was made. Prof.
D’Arcy Thompson also presented a few bottles and some
spirit to Dr. Donald. Mr. Bruce and Dr. Donald were
chosen by the Royal Geographical Society, and Mr. Leigh
Smith, to undertake the naturalist’s work, on board the
two whalers “‘ Balaena’”’ and ‘‘ Active,” which work they
voluntarily undertook.

On their return to Scotland in 1893 they presented their
collections, with the approval of Mr. Leigh Smith, to
the Museum of University College, Dundee, under the
care of Prof. D’Arcy Thompson, who forwarded the
Plankton to me for identification. It is regretable that I
have been unable to obtain certain notes on the collection,
which Mr. Bruce tells me both Dr. Donald and he handed
to Prof. D’Arcy Thompson on their return from the
Antarctic.

It appears that the collectors worked under very
considerable difficulties and in great discomfort on board
their vessels, greater even than might have been expected

*Trans. L’pool Biol, Soc., vol. XIL., p, 291,

ANTARCTIC PLANKTON. 189

‘on board a whaler. Hence the smallness of the collections.

Mr. Bruce mentions that the time they were actually in
the ice was from December 12th, 1892, to February
26th, 1893.

Referring to the mysterious presence of several fresh-
water animals in one of the bottles sent to me labelled
‘“ Antarctic,” Mr. Bruce thinks it probable that the bottle
in question was part of his Falkland Island fresh-water
collection. He also explains the occurrence of a number
of tropical species of Copepoda in another bottle (loc. cit.
p. 296) by the fact that they were collected by tow-net in
latitude 4° 02' N., longitude 23° 02’ W., as the missing
notes, supplied to the Dundee Museum with the collection,
showed.

Isaac C. THOMPSON.

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