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PROCEEDINGS |
| ‘.
TRANSACTRONS

IVERPOOL BIOLOGICAL SOCIETY.

-*
“We

WOU. TX:

SESSION 1894—95.

LIVERPOOL:

PRINTED BY T, Doss & Co., 229, BROWNLOW HILL.

1895,

i

1 ga rer i as, |
as ep ae eee ee ae ie oe,

574A, 0642

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§ Yaw. Aw |
CONTENTS.
I. PROCEEDINGS.
PAGE.
Office-bearers and Council, 1894—95 _ . : vali
Report of the Council . ‘re » Vill.
Summary of Proceedings at the Micsiinss 2 yy ees
Laws of the Society _ . : , : A Pane. air
List of Members . Kx
Librarian’s Report (with list of nies t ates 00.48
Treasurer’s Balance Sheet. ; Sy

IJ. TRANSACTIONS.
Opening Address :—‘“‘ Structure and Function.” By
FRANCIS GotcH, M.A., F.R.S., President : i
Highth Annual Report of the Liverpool Marine
Biology Committee and their Biological Station
at Port Erin. By Prof. HERDMAN,D.Sc.,F.R.S. 26

Description of Three Species of Anguillulide, ob-
served in Diseased Pseudo-bulbs of Tropical

Orchids. By Dr. J. G. de MAn : ace
Recent Additions to the Copepoda of Liverpool Bay.
By Isaac C. Tuompson, F.L.8., F.R.M.S. ~ oe OS

Report on the Investigations carried on in 1894 in
connection with the Lancashire Sea-Fisheries
Laboratory at University College, Liverpool.

By Professor W. A. Herpman, D.Sc., F.R.5.,
and Mr. ANDREW Scort, Fisheries Assistant. 104.

Note upon the yellow variety of Sarcodictyon caten-
ata, Forbes, with remarks upon the Genus and
its species. By W.A. Herpman, D.Sc.,F.R.S. 163

193212

iv. . CONTENTS.

Observations on the Tube-Forming Habits of Pan-

thalis oerstedt. By ARNoLD T. WATSON. . - ice
Observations upon the Pollen Tube. By A. J.
EWART, B.Sc. : : ; ; : .. £89

Notes on a Collection of Sponges from the West
Coast of, Portugal. By R. Hanirscu, Pu.D. 205
Notes on some points in the Structure of the Cerata
of -Dendronotus arborescens. By JosepH A.

Cheese, B-oC.s( Vick) 4 > a. 1a ’ . 220
Notes oh some Specimens of ‘Synapta inherens from
Port Hrn. By HERBERT C. CHADWICK . 285
Report on the Meduse of the L.M.B.C. District.
By Epwarp T. Browne, B.A. 243
Revision of the Amphipoda of the L.M.B.C. District.
By ALFRED O. WALKER, F.L.S. ‘ ‘2 . 287
A New Theory of Hearing. By C. HERBERT HuRstT,
PD, : 321

Report on Nemertines Pbepneert at eel ane in 11894
and 1895. By W. 1: Beaumont, bas . od04

PROCEEDINGS

OF THE

LIVERPOOL BIOLOGICAL SOCIETY.

OFFICE-BEARERS AND COUNCIL.

Ex-Presidents :

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

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

1891—92 T. J. MOORE, C.M.Z.S., A.L.S.

1892—93 ALFRED O. WALKER, J.P., F.L.S.

1893—94 JOHN NEWTON, M.R.C.S.

SESSION IX, 1894-95.

resident ;
Enon, F, GOTCH, M.A., F.RS,

Pice-Presidents :
Pror. W. A. HERDMAN, D.Sc., ¥F.R-S.
JOHN NEWTON, M.R.C.S.

Hon, Treasurer :
ISAAC ©. THOMPSON, F.L.S., F.R.M.S.

Hon. Librarian:
R. HANITSCH, Pu.D.

How. Secretary:
JOSEPH A. CLUBB, B.Sc. (Vrcr.).

Council :

H. C. BEASLEY.

H. O. FORBES, LL.D., F.Z.S.
Pror. R. J. HARVEY GIBSON, M.A.
W. J. HALLS.

J. SIBLEY HICKS, M.D., F.L.S.
ALFRED LEICESTER.

G. H. MORTON, F.G.S.
T. C. RYLEY.

W. -E. SHARP.

A. T. SMITH, Jun.

W. J. STEWART, B.A.
A. 0. WALKER, F.L.S.

REPORT of the COUNCIL.

Durine the Session 1894-95 there have been eight
ordinary meetings of the Society, held as heretofore at
University College, and two field meetings. The first
field meeting took place in May, to Hilbre Island.
Invitations were extended to members of the Geological
and Microscopical Societies of Liverpool, the Chester
Society of Natural Science, and the West Kirby Literary
and Scientific Society and a large and very successful
meeting was held. The second field meeting was to
Moreton and Leasowe Marshes in June, and was held
jointly with the Microscopical Society.

The communications made to the Society have been
representative of almost all branches of Biology, and
many interesting exhibits have been submitted at the
Meetings. |

The Library still continues to make satisfactory progress
as shown by the Librarian’s Report which follows. The
Council desire to record their appreciation of the services
rendered by Dr. Hanitsch, who has acted as Hon. Librarian
for some years, and under whose fostering care the Society’s
Library has made such satisfactory progress. Dr.
Hanitsch’s work in arranging exchanges with other scien-
tific societies has been most valuable.

The Treasurer’s usual statement and Balance Sheet are
appended. No alterations have been made in the Laws
of the Society during the past session. The membership —
has been well sustained, twenty new members having
been elected during the past session.

The members at present on the roll are as follows :—

Honorary Members ......... 7
Ordinary Members 222... 2. 66
Student Weniberse sem 25

SUMMARY of PROCEEDINGS at the MEETINGS.

The first meeting of the ninth session was held at
University College on Friday, 12th October, 1894; Prof.
Gotch, F.R.S., President, in the chair.

1. The Report of the Council on the Session. 1893-94 (see
“‘ Proceedings,” Vol. VIII., p. vil.) was read and
adopted.

2. The Treasurer’s Balance Sheet for the Session 1893-94
(see ‘‘ Proceedings,’ Vol. VIII., p. xxix.) was
submitted and approved.

3. The Librarian’s Report (see ‘‘ Proceedings,” Vol. VIII.,
p- XXv.) was submitted and approved.

4. The following Office-bearers and Council for the ensuing
Session were elected :— Vice-Presidents, Prof. W. A.
feraman, 1).Sc., H.R.S., John Newton, M.R.C.8.;
frome ereasurer, |. C.'Thompson, F’.L.5., F.R.M.S.;
fron, Mnbrarian, R. Hanitsch, Ph.D. ;.Hon. Secre-
tary, Joseph A. Clubb, B.Sc. (Vict.) ; Council, H.
Seeeeasicy, 4. O. Forbes, LL.D., F.Z.8., R. J.
fiavey Gibson, M-A.; F.I:8., W. J. Halls, J.
Sibley Hicks, M.D., Alfred Leicester, G. H. Morton,
ieee, 1. C. Ryley, W. E. Sharp, A. T. Smith,
gun., W.J. Stewart, B.A.,and A. O. Walker, F.L.S.

5. Prof. R. J. Harvey Gibson exhibited some recent
additions to the Herbarium of the Botanical Depavrt-
ment of University College.

6. Mr. Joseph A. Clubb, B.Sc., exhibited with remarks a
series of trays illustrating Insect metamorphoses
from the Bootle Free Public Museum.

7. Dr. H. O. Forbes exhibited a living specimen of the
African Mud-fish (Protopterus annectens) with re-

x PROCEEDINGS LIVERPOOL BIOLOGICAL SOCIETY.

marks. It created great interest and additional
observations were made by Prof. Herdman and
the President.

8. The President (Prof. Gotch, F.R.S.), delivered the
Presidential Address, entitled ‘‘ Structure and
Function” (see ‘‘ Transactions,” p. 1). A vote of
thanks, proposed by Dr. Hurst, seconded by Mr.
Alf. O. Walker, was carried with acclamation.

The second meeting of the Society was held at University
College on Friday, November 9th, 1894, Prof. Gotch,
President, in the chair.

1. Mr. G. H. Morton, F.G.S8., exhibited a list of subscribers
to an old work by Mr. John Price, on the “‘ Fauna
of the Birkenhead Shore,” and also a work by Mr,
Isaac Byerley, dated 1852, on the “ Fauna of
Liverpool Bay and its neighbourhood.”

2. Dr. Grossmann exhibited some beans containing an
insect larva, which when heat was applied caused
the beans to move.

3. Mr. I. C. Thompson, F.L.S., described some new
species of Copepoda and exhibited some drawings
of .the same made by Mr. Andrew Scott (see
“Transactions,” p. 95).

4. Prof. Herdman, F.R.S., gave the Annual Report of
the Marine Biology Committee and the Port Erin
Biological Station (see “‘ Transactions,” p. 26, and
Pls. I. and II.). The report was illustrated by
lantern slides and specimens.

5. Rev. T. 8. Lea exhibited a series of lantern slides
prepared from micro-photographs of marine alge,
taken by himself at Port Erin.

6. A paper on three new species of Nematodes found in
Orchids, by F'. G. de Man, Ph.D., was submitted (see
“Transactions,” p. 76. and Pls, II1., IV. and V.).

7

. SUMMARY OF PROCEEDINGS AT MERTINGS. Xl.

_ The third meeting of the Society was held at University
College on Friday, December 14th, 1894, Prof. Gotch,
President, in the chair.

1. A paper by Mr. A. J. Ewart, B.Sc., entitled ‘“‘ Observa-
tions on the Pollen-tube,’’ was submitted by Prof.
Harvey Gibson (see ‘“‘ Transations,”’ p. 189, Pl. XI.).

2. Prof. Herdman, F.R.S., gave a short note on a species
of Sarcodictyon from Port Erin (see ‘‘ Transactions,”
patos, and Pl. ViIT.).

3. Dr. Hurst read a paper entitled ‘‘A new Theory of
Hearing’”’ (see ‘‘ Transactions,’”’ p. 321). By in-
vitation of the Council a number of the members
of the Liverpool Physical Society were present,
and an animated discussion took place, in which
remarks were made by Prof. Lodge, Dr. Howard,
Dr. Paul, the President and others.

The fourth meeting of the Society was held at University
College on Friday, January 11th, 1895, Prof. Gotch, F.R.5.,
President, in the chair.

1. Dr. Karl Grossmann gave an account of some recent
excavations at Pompeii, and described some of the
domestic and sanitary arrangements existing before
the great eruption. Dr. Newton added some
further interesting details.

mr, HH, O. Forbes and Prof. Herdman, F.R.S., des-
cribed some of the anatomical peculiarities of the
tail of a Siamese Cat belonging to Mr. Richard D.
Holt. Mr. Ridley of the Botanical Gardens, Singa-
pore, added some further remarks on the habits of
the animals. The skeleton and stuffed specimens
were exhibited.

8. Mr. H. C. Chadwick read a short paper on the structure
of a species of Synapta from Port Erin (see

- Xil. PROCEEDINGS LIVERPOOL BIOLOGICAL SOCIETY.

“Transactions,” p. 235, and Pls. XVI. and XVIL.).
4. Mr. Arnold T. Watson gave a paper on the tube forming
habits of Panthalis oerstedi (see ‘‘ Transactions,”
p. 169, and Pls. IX. and X.).. The paper was
illustrated by lantern slides. |

The fifth meeting of the Society was held at University
College on Friday, February 8th, 1895, Dr. Newton, Vice-
President, in the chair.

1. Mr. W. E. Sharp gave an interesting note on Sete
abrata var. swbrotundata, and exhibited specimens
of the same.

2. Mr. Andrew Scott briefly laid before the Society the
Annual Report on the investigations carried on in
the Fisheries Laboratory, University College, by
Prof. Herdman and himself (see ‘‘ Transactions,”
p. 104).

3. Dr. Karl Grossmann gave an interesting account of a
recent visit made by himself and two friends to the
Faroe Islands. He vividly described the scenery
and mentioned some interesting facts concerning
the Fauna and Flora of the Islands. The Vice-
President, the Rev. T. S. Lea and others added
some remarks. An excellent series of photo-lantern
slides beautifully illustrated the lecture.

The sixth meeting of the Society was held at University
College on Friday, March 8th, 1895. In the absence of
the President, Mr. I. C. Thompson took the chair.

1. Prof. Harvey Gibson exhibited and described some
models and microscopic preparations recently added
to the Botanical Department of University College.

2. Mr. G. H. Morton in a short note on the supposed
occurrence of Fusulina at Penmon, Anglesea,

SUMMARY OF PROCEEDINGS AT MEETINGS. X11.

expressed his belief that this fossil foraminifer did
not really occur there. He exhibited specimens of
the real fossil, which created great interest.

8. Some additional notes on the LucERNARIANS and
on the Nemertina of Port Erin, byW. I. Beaumont,
were communicated by the Hon. Secretary (see
Eliraneactions,’ p.-354).:—

4, Dr. Hanitsch read a short paper on a collection of
fresh-water sponges. He described some anatomical
peculiarities, and added some general remarks on
the geographical distribution of fresh-water sponges.

5. Mr. Andrew Scott brought forward a record of additions
to the Fauna of the L.M.B.C. District, and exhibited
drawings of some new species of Copepoda.

6. The Report of the Mepusz of the L.M.B.C. District
by Mr. E. T. Browne (see “ Transactions,”’ p. 248).

_{Communicated by the Hon. Secretary. ]

The seventh meeting of the Society was held at University
College on Friday, April 5th, 1895, Prof. Herdman, Vice-
President, in the chair.

1. Miss F. Phillips exhibited some microscopic prepara-
tions of mites, and some drawings of the same.

2. Mr. Alf. O. Walker, F.L.8., laid before the Society a
revision of the AmpHipopaA of the L.M.B.C. District
(see ‘‘ Transactions,” p. 287, and Pls. XVIII. and
XIX.).

8. Mr. Joseph A. Clubb, B.Sc., read a paper on the
relations of the hver and the cerata in Dendronotus
arborescens (see ‘‘ Transactions,” p. 220, and Pl.

XIV.).

The eighth meeting of the Society was held at University
College on Friday, May 17th, 1895, Prof. Gotch, President,
in the chair.

XIV. PROCEEDINGS LIVERPOOL BIOLOGICAL SOCIETY.

1. On the proposition of the President, seconded by Prof.

oo) Mit:

5. M

BS

Herdman, the best thanks of the Society were
accorded to Dr. Hanitsch, who was leaving England,
for his services as Hon. Librarian. Dr. Hanitsch
suitably responded.

r. H. O. Forbes exhibited some specimens of Malap-

terurus electricus, in which the electric organ had
been dissected out and was exposed to view. The
President added some interesting details of the
minute anatomy, and referred to the exhibition of
very excellent microscopic preparations made by
Mr. Macdonald, B.A.

H. C. Beasley exhibited with remarks some stone
arrow-heads from Northern Chili.

. Hanitsch briefly described a collection of sponges

from the West Coast of Portugal (see “‘ Tran-
sactions,”’ p. 205, and Pls. XII. and XII).

. Joseph A. Clubb, B.Sc., gave a note on the inner-

vation of the cerata of Dendronotus arborescens
(see ‘‘ Transactions,” p. 226, and Pl. XV.).

. H. O. Forbes exhibited specimens of the remains of

Aphanapteryx hawkinst and Pal@ocorax morirum
from.the Chatham Islands, and gave an interesting
account of the distribution of life in the Southern
Hemisphere, advancing certain arguments therefrom

in favour of the previous existence of an extensive

Antarctic Continent. An interesting discussion
followed, but it was felt that the field opened up
by Dr. Forbes’ remarks was too extensive to do
justice to that evening, and the hope was expressed
that Dr. Forbes would, at a subsequent meeting
next session, give the Society an opportunity of
again hearing his views.

7. At the close of the meeting Prof. Herdman congratu-

SUMMARY OF PROCEEDINGS AT MERTINGS. XV.

lated, in the name of the Society, the President
(Prof. Gotch, F.R.S.), on his recent appointment
to the Waynflete Professorship of Physiology at
Oxford, and expressed the hope that the Society
might still be sometimes favoured by his presence.
Prof. Gotch in returning thanks for the good
wishes expressed on his behalf, stated his intention
of continuing his connection with the Biological
Society.

The first field meeting was held at Hilbre Island at the
mouth of the Dee on May 25th. By invitation the party
was joined by members of the Geological and Micros-
copical Societies of Liverpool; the Chester Society of
Natural Science, and the West Kirby Literary and
Scientific Societies. A very large and successful meeting
was held.

The second field meeting of the session took place on
June 8th at Moreton and Leasowe Marshes. On this
occasion a joint meeting was held with the Liverpool
Microscopical Society. After the ponds and ditches in the
neighbourhood of the Lighthouse had been explored, the
party sat down to tea. After tea a short business meeting
was held, Prof. Herdman, Vice-President, in the chair.
On the motion of Prof. Herdman, seconded by Mr. A. T.
Smith, Jun., Prof. Harvey Gibson, M.A., was elected
President for the coming session.

LAWS of the LIVERPOOL BIOLOGICAL
SOCIETY.

I.—The name of the Society shall be the ‘‘ LIVERPOOL
BroLoaicaL Society,” and its object the advancement of
Biological Science.

JI.—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.

III.—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.

ViI.—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

oS

LAWS. XVIL

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 ex-

clude. -

VItI.—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.

1X.—E very person so elected shall within one calendar

month after the date of such election pay an Kntrance Fee
of Half a Guinea and an Annual Subscription of One

>< LIVERPOOL BIOLOGICAL SOCIETY.

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-
scriptions 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 Subseription 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 the 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 am
writing to the Secretary, but the Member so resigning shall
be liable for the payment of his Annual Subscription, and
all arrears up to the 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

LAWS. a -X1X.

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, ar any ten Members
can convene a Special General Meeting, to be called within
fourteen days, by giving notice in writing to the Secretary,
and stating the object of the desired Meeting. The circu-
lar 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,

oy AX. ; LIVERPOOL BIOLOGICAL SOCIETY.

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-LAW.

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.

kA

LIST of MEMBERS of the LIVERPOOL

ELECTED.

1890
1886

1890
1886

1888
1894
1889
1894
1887
1886
1886

1890
1891
1889

1886
1890
1894

1891

BIOLOGICAL SOCIETY.

SHSSION 1894-95.

A. ORDINARY MEMBERS.

(Life Members are marked with an asterisk. )

Assheton, R., M.A., Owens College, Manchester

ipamkes rot, W. Mitchell, M.D., F!R.C.8., 28,
Rodney-street |

feet. A. ti, B.A. LIB: F.L.8., The
Laurels, Wormley, Herts

iaerom, Prot. Alexander, M.B., M.R.C.S8., 31,
Rodney-street

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

Boyce, Prof., University College, Liverpool

Brown, Prof. J. Campbell, 27, Abercromby-square

Browne, A. H., M.D., 67, Catherine-street

Caine, Nathaniel, Spital, Bromborough

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

Clubb, J. A., B.Sc., Hon: SEcRETARY, Free
Public Museum, Liverpool

Davies, D., 55, Berkley-street

Dismore, Miss, 65, Shrewsbury-road, Oxton

Dwerryhouse, A. R., 8, Livingstone Avenue,
Sefton Park

Ellis, J. W., M.B. (Vic.), F.E.S., 18, Rodney-st.

Ewart, A. J., B.Sc., University College, Liverpool

Forbes, H.O., LU.D., F.Z.8., Free Public Museum,
Liverpool

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

XX. LIVERPOOL BIOLOGICAL SOCIETY

1886 Glynn, Prof. T. R., M.D., F.R.C.P., 62, Redmeg =a
street

1886 Gibson, Prof. R. J. Harvey, M.A., F..8., Univer- ~
sity College |

1891 Gotch, Prof. F., F.R.S., PRastpEnt, Oxford :

1894 Gould, J., Littledale-road, Egremont |

1894 Greening, Linneus, F.L.8., 5, Wilson Patten-
street, Warrington

1894 Grossmann, Karl, M.D. , Rodney- street, i

1886 Halls, W. J., 35, Lord-street

1887 Hanitsch, R., Ph.D., Raffles Museum, Singapore |

1887 Healey, Ghee ee Galcialel Gateacre q

1886 Herdman, Prof. W. A., D.Sce., FBS.) Viena ae
PRESIDENT, University Colles

1893 Herdman, Mrs., B.Sc., 32, Bentley-road, Liven :

[Sol lickssd. csibley. M.D., 2, Erskine-street :

1894 Hickson, Prof. S$. J., Owens College, Manchester

1888 *Hurst, C. H., Ph.D., Owens College, Manchester

1886 Jones, Charles W., Field House, Prince Altred- —
road, Wavertree ;

1894 Jones, Charles EK. Elpie, Prenton-rd. W., B’head

1894 lea, Rev. T. 8., 3, Wellington Fields, Wavertree

1886 Leicester, Alfred, 30, Weld-road, Birkdale

1886 Lomas, J., Assoc.N.8.8., Salen, Amery Grove,
Birkenhead

1893 Macdonald, J. S., B.A., Physiological Lab. Univ. |
College, Liverpool

1888 Melly, W. R., 90, Chatham-street :

1886 McMillan allen S., F.L.8., Brook-road, Maghull =

1886 Morton, G. H., F.G.S., 209, Hdge-lane, H.

1888 Newton, John, M.R.C.S., Vicz-PresipEent, 44,
Rodney-street

1887 Narramore, W., F.L.S., 5, Geneva-road, Elm Park

1894 Paterson, Prof., University College, Liverpool

LIST OF MEMBERS. XX.

| 1894 Paul, Prof. F. T., Rodney-street, Liverpool

1891 Phillips, Miss F., 3, Green-lawn, Rock Ferry

1892 Phillips, HK. J. M., L.D.S., M.R.C.S., Rodney-st.

1886 *Poole, Sir James, J.P., Abercromby Square

1890 Rathbone, Miss May, Backwood, Neston

1890 Roberts, Lesle, M.D., 31, Rodney-street

1887 Robertson, Helenus R., Springhill, Church-road,
Wavertree

1887 Ryley, Thomas C., 10, Waverley-road

1892 Sephton, Rev. J., M.A., 90, Huskisson-street

1891 Sharp, W. E., The Woodlands, Ledsham, Chester

1886 Smith, Andrew T., Jun., 13, Bentley-road, Prince’s
Park

1889 Stewart, W. J., B.A., Magistrates’ Court, Dale-st.

1893 ‘Tate, Francis, F.C.S., Hackins Hey, Liverpool

foo Phompson, Isaac C., F.L.8., F.R.M.8., Hon.
TREASURER, Woodstock, Waverley-road

1889 Thornely, Miss L. R., Baycliff, Woolton Hull

1888 Toll, J. M., Kirby Park, Kirby

1886 Walker, Alfred O., J.P., F.L.8., Colwyn Bay

1889 Williams, Miss Leonora, Hill Top, Bradfield, nr.
Sheffield

1891 Wiglesworth, J., M.D., County Asylum, Rainhill

1891 Wood, G. W., F.1.C., Riggindale-road, Streatham,
London

1892 Weiss, Prof., Owens College, Manchester

1892 Young, T. F., M.D., 12, Merton-road, Bootle

B. STuDENT MEMBERS.

_ Armstrong, Miss A., 26, Trinity-road, Bootle
Bates, H., University College, Liverpool
Crowther, H. P., University College, Liverpool
Chadwick, H. C., Free Museum, Bootle
Christophers, S. R., 10, Lily-road, Fairfield

XX1y.- LIVERPOOL BIOLOGICAL SOCIETY.

Depree, 8. 8., 8, Morley-road, Southport

Dickinson, T., 3, Clark-street, Prince’s Park

Dumergue, A. F.,7, Montpellier-terrace, Up. Parliament-st.
Dutton, J. E., Kings-street, Rock Ferry

Ewart, R. J., University College, Liverpool

Hannah, J. H. W., 8, Allington-street, Aigburth-road
Hamilton, D., 16, Whitefriars, Chester

Harvey, EH. J. W., 5, Cairns-street, Liverpool

Hawkes, A. HE. W., 22, Abercromby-square, Liverpool
Hay, John, 92, Bridge-street, Birkenhead |
Henderson, W. 8., B.Sc., 2, Holly-road, Fairfield
Hurter, D. G., Holly Lodge, Cressington

Linton, §. F., St. Pauls Vicarage, Clifton-road, Birkenhead
Lowe, O. W. A., 4, Wexford-road, Oxton

Quinby, F. G., 11, Belvidere-road, Liverpool

Simpson, A. Hope, Annandale, Sefton Park —

Scott, A., University College, Liverpool

Sykes, A. B., Manor House, Formby

Warham, Miss A. E., B.Sc., 70, North-st. St. Andrews N.B.
Willmer, Miss J. H., 20, Lorne-road, Oxton, Birkenhead

C. HonoRARY MEMBERS.

H.8.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. E., University, Jena

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

REPORT of the LIBRARIAN.

Our Society has arranged four additional exchanges of
publications since the last Report, making in all seventy-
six societies and institutions.

The following list gives the titles of the exchanges and
donations received during the session :—

Bi
18.

Allgemeine Fischerei-Zeitung. XIX.,10-—26; XX., 1—9.

Amsterdam ; Verhandelingen der K. Akademie van Wetenschappen.
(S. 2), Vols. I—III.

Annaes de Sciencias Naturaes. I., Nos. 1, 2, 3, and 4. II., No. i.

Annaes de Sciencias Naturaes publica dos por Augusto Nobre. L.,
Wow; i., INo;: 2.

Archives Néerlandaises des Sciences exactes et naturelles. XXVII.,

3—5; XXVIII., 1—5; XXIX., 1.

Australian Museum, The; Report of the Trustees for the year 1893.

Bergens Museums Aarbog for 1893.

Bergens Museum. V. On the development and structure of the
Whale. PartI. By G. Guldberg and Fridtjof Nansen.

Berlin, Math. u. naturw. Mittheilungen d. k. preuss. Akademied.,
Wissenschaften, April, July, October, November, 1894; January
February, 1895.

Berlin, Sitzungsberichte d. k. press. Akademie d. Wissenschaften.
1894, Nos. 1—58.

Bihang till K. Svenska Vetenskaps—Akademiens Handlingar. XIX.,
Nos. 3 and 4,

Bologna, Memorie della R. Accademia delle Scienze dell’ Istituto.
te. 3), ITI.

Bordeaux, Procés—Verbaux de la Société Linnéenne. Vol. XLVI.
(1893).

British Naturalist. Nos. 5—8.

Canada ; Bulletin of the Natural History Society of New Brunswick.
XII.

Canada, Proceedings and Transactions of the Nova Scotian Institute
of Science. Session 1892—93.

Canadian Institute ; Seventh Annual Report. Session 1893-94.

Canadian Institute ; Transactions. IV., No. 1.

XXVI1.

LIVERPOOL BIOLOGICAL SOCIETY.

Chili; Actes de la Société Scientifique. Vol. II., No. 4; III., Nos.
3—5; IV., Nos. 1—4.

Chili; Revista Chilena de Hijiene. Vol. I., Nos. 1 and 2.

Christiania, Videnskabs—Selskabs Forhandlingen for 1893.

Denmark, Oversigt over det K. Danske Vidensk. Selskabs Forhand-
linger, 1893, No. 3; 1894, Nos. 1 and 2.

Ergebnisse der Beobachtungsstationen an den deutschen Kiisten, 1893
Nos. 1—6.

Frankfurt a. M.; Bericht itber die Senckenbergische naturforschende.
Gesellschaft, 1894.

Gottingen ; Nachrichten d. k. Gesellschaft d. Wissenschaften, 1894,
Nos. 1—4; 1895, No. 1.

Japan, Teena of the College of Science, bapa University. Vol.
VI., part 4: VIL., part 1—3; VHL., part 1.

Kjobenhayn ; Videnskabelige Meddelelser fra der naturhist, 1894.

Lancashire and Cheshire Entomological Society ; Seventeenth Annual
Report, 1893.

Leipzig ; Berichte d. k. Gesellschaft d. Wissenschaften, 1894, II.

Madras Government Museum. Bulletin Nos. 1—3. |

Manchester Microscopical Society. Trausactions and Annual Report,
1893.

Marine Biological Association of the United Kingdom. Journal (N.S.),
III., Nos. 3 and 4.

Marine Zoology and Microscopy. Journal, 1—3 and 4.

Meriden Scientific Association. Vol. V.

Montpellier, Académie des Sciences en Lettres. Mémoires de la
section des Sciences. (S. 2), I, Nos. 1, 3 and 4.

Moscou, Bulletin de la Société Imperiale des Naturalistes, 1893, No.

4; 1894, 1—4.

Muséum d’Histoire naturelle. Bulletin Année 1895, Nos. 1 and 2. .

Nancy, Bulletin de la Société des Sciences. (S. 2), XIII..

Napoli, Rendiconto dell’ Accademia delle Scienze fisiche e matematiche.
Vililils, Noss 612 (Si-3), I.,, Noss 1-2:

Natuurkundig Tijdschrift voor Nederlandsch—-Indie. LIII.

New Zealand Institute, Transactions and Proceedings of the, 1893.
Vol. XXVI. .
Revue Biologique du Nord de la France. V., Nos. 3—12, VI., VIL,
Nos. 1—38.
Rheinlande, Verhandlungen d. naturh. Vereins d. preuss. Vol. L.,

INow2);L1.5 Now
Royal Boiss of Physicians, Edinburgh, Reports from the ma
of the. Vel. ¥. i 003 .

LIBRARIAN’S REPORT. XXVii.

Royal Dublin Society, Transactions of the. Vol. IV., No. 14; Vol.
V., Nos. 1—4.
Royal Dublin Society, Proceedings of the. Vol. VII., part 5;
Sev, part | and 2.
Royal Microscopical Society, Journal of the. 1892, 1893, 1894,
Nos. 1—6.
Royal Physical Society Edinburgh, Proceedings of the. Sessions
1892-93 and 1893-94.
Scottish Fishery Board, Annual Reports, XI. and XII.
Stavanger Museum. Aarsberetning for 1893. —
St. Pétersbourg. Bulletin de l Académie Impeériale des Sciences (N.S.),
Vol. 1V., Nos. 1 and 12 (S. 5), Vol. I., 1—3 (S. 5), Vol. II., 1—3.
The Naturalist. Monthly Journal of Natural History for the North of
England. May—December, 1894; January—April, 1895.
Torino, Bolletino dei Musei di Zoologia ed Anatomia comparata delle
R. Universita. Vol. IX.
United States, Bulletin of the Fish Commission. Vols. XI., XII.
and XIII.
United States, Commission of Fish and Fisheries. Report of the
Commissioner. 1889—91 and XVIII.
United States, Proceedings of the National Museum. Nos. 975—
979, Vol. XVII., Nos. 999-1009.
United States, Bulletin of the National Museum. No. 438.
United States, Report of the National Museum for 1892, pp. 245—
368, 461—493.
United States, Annual Reports of the Board of Regents of the Smith-
sonian Institution, 1892 and 1893.
United States, Smithsonian Institution. Report of the National
Museum. 1891 and 1892.
United States, Smithsonian Institution, Proceedings ‘of National
National Museum. Nos. 1011—1017.
United States, Annual Report, Museum of Comparative Zoology,
Harvard College, 1893-94.
United States, Bulletin of the Museum of Comparative Zoology,
Harvard College. XXV-., Nos. 7—11; XXVI., No. 1.
United States, Proceedings of the Academy of Natural Sciences of
Philadelphia, 1894, parts 1—2.
United States, Proceedings of the Boston Society of Natural History.
Vol. XXVI.
United States, Transactions of the Academy of Science of St. Louis,
VI., Nos. 9—17.
United States, Tufts College Studies. No. 2 and 3,

XXVIIl. LIVERPOOL BIOLOGICAL SOCIETY.

68.
69.

70.

(ale

89.

90.

Victoria, Proceedings of the Royal Society of. Vol. I. (N.S.). .

Wien, Verhandlungen der k. zoologisch-—botanischen Gesellschaft.
XLIV., Nos. 1—4; XLV., 1—38. ‘:

Wien, Annalen des K. K. Naturhistorischen Hofmuseums. VIII.
Nos. 2—4.

Wissenschaftliche Meeresuntersuchungen d. Kommission zur wiss.
Untersuchung der Deutschen Meere in Kiel u. der Biologischen
Anstalt auf Helgoland. (N. F.), I., No. 1.

Ziivich, Vierteljahrschrift d. naturforschenden Gesellschaft. XXXIX.,
Ness? 4; Nise Niow It

Zeitschrift fiir Fischerei. I., 6; II., 1—4.

Lilljeborg, V. Synopsis Crustaceorum Svecicorum ordinis Branchiopo-
dorum et subordinis Phyllopodorum, 1877.

Théel, Hj. Note sur quelques Holothuries des mers de la Nouvelle
Zemble, 1877.

Tullberg, T. Ueber die Byssus des Mytilus edulis, 1877.

Wittrock, V. B. On the development and systematic arrangement of
the Pithophoracee, a new order of Alg, 1877.

Glas, O. Essai sur la Société Royale des Sciences d’ Upsal et ses
rapports avec l’université d’ Upsal, 1877. ?

Kjellman, F. R. Ueber die Algenvegetation des Murmanschen Meeres
an der Westkiiste von Nowaja Semlja und Wajgatsch, 1877.

Fristedt, R. F. Ioannis Franckenii Botanologica nune primum
edita, praefatione historica, annotationibus criticis, nomenclatura
Linneana illustrata, 1877.

Fries, Th. M. Polyblastize Scandinavice, 1877.

Almén, Aug. Analyse des Fleisches einiger Fische, 1877.

Lundstrom, A. N. Kritische Bemerkungen iiber die Weiden Nowaja
Semljas und ihren genetischen Zusammenhang, 1877.

Lindman, C. A. M. Om postflorationen och dess betydelse sasom
skyddsmedel for fruktanlaget, 1884.

Wirén, A. Om cirkulations—och digestions—organ en hos Annelider
af familjerna Ampharetide, Terebellide och Amphictenide, 1885.
Fristedt, K. Bidrag till kannedomen om de vid Sveriges vestra kust

lefvande Spongiz, 1885.

Appellof, A. Japanska Cephalopoder, 1886.

Henning, E. Agronomiskt Vaxtfysiognomiska Studier i Jemtland,
1889.

Tedin, H. Bidrag till kinnedomen om primiara barken hos vedartade
Dikotyler, dess anatomi och dess funktioner, 1891.

Lonnberg, E. Anatomische Studien iiber Skandinavische Cestoden,
1891.

oT

- 92.

100.
101.
102.
103.
104.

105.

106.
107.

108.

109.
110.
Lu.
112.

113,

114,

LIBRARIAN'S REPORT. Ode e

Schott, H. Zur Systematik und Verbreitung palearctischer Collem-
bola, 1893.

Wetterdal, H. Bidrag till kannedomen om Bakteriehalten i Vatten-
dragen invid Stockholm, 1894.

Lilljeborg, W. Bidrag till kannedomen om Pterycombus Brama B,
Fries, en Fisk af Makrillfiskarnes familj.

Tilljeborg, W. Bidrag till kannedomen om underfamiljen Lysianas-
sina inom underordningen Amphipoda kraftdjuren.

Lilljeborg, W. Ofversigt af de inom Skandinavien (Sverige och
Norrige) antraffade Hvalartade Daggdjur (Cetacea), 1861.

Fries, E. Epicrisis generis Hieraciorum, 1862.

Fristedt, R. F. Studier i Allman Pharmakognosi.

Smitt, F. A. Bidrag till kannedomen om Hafs Bryozoernas utveckling.

Friies, O. R. Om Svampbildningar pa menniskans hud och deraf
fororskade, sjukdomar, 1867.

Fristedt, R. F. Preparata pharmacotechnica, 1866.

Lilljeborg, W. Bidrag till kannedomen om de inom Sverige och
Norrige forekommande Crustaceer af Isopodernas underordning och
Tanaidernas familj.

Ekecrantz, W. Bidrag till kinnedomen om de i menniskans tarmkanal
forekommande Infusorier, 1869.

Bovallius, C. Om Balanidernas utveckling, 1875.

‘Upsala Universitets Arsskrift, 1875.

Hogman, S. Jemforande framstallning af skelettbyggnaden hos
Colymbus och Podiceps, 1873.

Arnell, H. W. Om vegetationens utveckling i Sverige aren, 1873—75.

Aurivillius, C. W. 8. Bidrag till kannedomen om Krustaceer, som
lefva hos Mollusker och Tunikater, 1883.

Forsstrand, C. Det arktiska hafsomradets djurgeografiska begrainsing
med ledning af shalkraftornas (crustacea malacostraca) utbredning,
1886.

Bayer, S. Bidrag till kannedomen om Bakterierna i Menniskans
Tarmkanal, 1886.

Juel, H.O. Beitraige zur Anatomie der Trematodengattung Apoblema
(Dujard.), 1889.

Stenstrom, K. O. E. Yarmlandska Archhievacier anteckningar till
Skandinaviens Hieracium-Flora, 1890.

Skarman, J. A.O. Om Salixvegetationen i Klarelfvens Floddal, 1892.

Hedland, J. F. Kritische Bemerkungen iiber einige Arten~ der
Flechtengattungen Lecanora, Ach., Lecidea, Ach., und Micarea,
Fr., 1892.

Hedenius, P. Om Upptackten af Blodomloppet, 1892,

.

EER:

115.
116.
117.
118.
119.

120.

17a.
122.

123.

“wt ==) ee
’ '

LIVERPOOL BIOLOGICAL SOCIETY.

Sundberg, C. J. G. Undersdékningar ofver mojlicheten af Mikrobers
intrangande genom den oskadade tarmslemhinnans yta, 1892.

Elfstrand, M. Hieracia alpina aus den Hochgebirgsgegenden. des
mittleren Skandinaviens, 1898.

Jagerskidld, L. A. Bidrag till kinnedomen om Nematoderna, 1893.

Ekholm, K. Studier ofver Kolonbakterien, 1893.

Starback, K. Studier; Ellias Fries’ Svamp-Herbarium. I. ‘‘Sphe-
riacese imperfecte cognite,” 1894.

Eliasson, A. G. Om Sekundira, Anatomiska Forandringar inom
Fanerogamernas Florala Region, 1894.

Levander, K. M. Beitrage zur Kenntniss einiger Ciliaten, 1894.

Segerstedt, P. Studier ofver buskartade stammars skyddsvafnader,
1894. | ;

Fries, Th. M. Bidrag till en lefnadsteckning ofver Carl von Linné,

(Nos. 74—123 have been presented by the Royal University of Upsala).

124,

125.

126.

133.

Il genere Ankylostomum, Dubini.--Osservazioni sul Solenophorus
megalocephalus.--I Distomi dei Rettilii—Notizie elmintologiche.
By Michele Stossich. Presented by the author.

Report on the Danish Biological Station to the Home Department.
IV. (1893). By C. G. Joh. Petersen. Presented by the author.
Sur les Nerfs de l’antenne et les Organes chordotonaux chez les
Fonrmis. Sur le systeme glandulaire des Fourmis.—Pedodera des
glands pharyngiennes de Formica rufa, L.—Sur la morphologie du
squelette des segments post thoraciques chez les Myrmicides.—Sur
V anatomie du pétiole de Myrmica rubra, L.—Transformation
artificielle en Gypse du Calcaire friable des Fossiles des Sables de
Bracheux. Par Charles Janet. Presented by the author.

A Monograph of the Mycetozoa, by Arthur Lister. Presented by the
British Museum.

Regles de la Nomenclature des étres organisés. Presented by the
Zoological Society of France.

. Der Elbelachs. Eine biologisch—anatomische Studie. By Prof. Ant.

Fritsch. Presented by the author.

Untersuchungen iiber die Fauna der Gewasser Bohmens. No. IV.
By Prof. Ant. Fritsch and Dr. V. Vavra. Presented by the authors,

Contributions towards a knowledge of the anatomy of the Genus
Selaginella, Spr. By Prof. R. J. Harvey Gibson, Presented by
the author.

Note on the Diagnostic Characters of the subgenera and species of
Selaginella, Spr. By Prof. R. J. Harvey Gibson. Presented by
the author.

Uber die Metamorphose der Siisswasser—Ostracoden. By Prof. C,
Claus, Presented by the author.

.
wow elm Tl

LIBRARIAN'S REPORT. ~ XXX1.

F _ 134, Resultats des Campagnes Scientifiques accomplies sur son yacht par
Albert Ie, Prince Souverain de Monaco. Fasc. VII., Brachyures
et Anomoures. Par A. Milne—Edwards et E. L. Bouvier. Pre-
sented by H.S.H. the Prince of Monaco.

135. Museums of the Past, the Present, and the Future, particularly those

. of Liverpool. By G. H. Morton, F.G.S. Presented by the author.

136. Bericht tiber die Feier des 60. Geburtstages von Ernst Hieckel.
Presented by the Committee.

-List of Societies, etc., with which publications are
exchanged.

AmsrerpDAM.—Koninklijke Akadamie van Wettenschappen.

Koninklijke Zoologisch Genootschap Natura Artis Magistra.

BAttTImorE.—Johns Hopkins University.
Barayia.—Koninklijke Natuurkundig Vereeniging in Ned. Indie.
Bercen.—Museum.
Beriry.—Konigl. Akadémie der Wissenschaften.
- Deutscher Fischerei-Vereins.
BirMINGHAM.——Philosophical Society.
Botonea.—Accademia delle Scienze.
Bonn.—Naturhistorischer verein des Preussischen Rheinlande und West-
falens.
Borpeaux.—Société Linnéenne.
Bosron.—Society of Natural History.
Brussevts.—Academie Royale des Sciences, etc., et Belgique.
CAMBRIDGE.—Morphological Laboratories.
Campriper, MAss.—Museum of Comparative Zoology of Harvard College.
CuRIstIANIA.—-Videnskabs-Selskabet.
Dusiin.—Royal Dublin Society.
EprnsurGcu.—Royal Society.

Royal Physical Society.

Royal College of Physicians.

Fishery Board for Scotland.
Frankrurt.—Senckenbergische Naturforschende Gesellschaft.
Frerpure.—Naturforschende Gesellschaft
~ Gunuvn.—Société de Physique et d'Histoire Naturelle.
Ginssen.-—Oberhessische Gesellschaft fiir Natur und Heilkunde,
GLAscow,—Natural History Society.

XXX. ' LIVERPOOL BIOLOGICAL SOCIETY.

Gortincrn.—Konigl. Gesellschaft der Wissenschaften.
Hauirax.—Nova Scotian Institute of Natural Science.
HARLEM.—Musée Teyler.

Société Hollandaise des Sciences.
Kre..—Naturwissenschaftlichen vereins fur Schleswig—Holstein.
Kommission fur der Untersuchung der Deutschen Meere.
KsopenHAyn.—Naturhistoriske Forening.

Danish Biological Station, (C. G. John Petersen).

Kongelige Danske Videnskabernes Selskab.
Lreps.—Yorkshire Naturalists Union.
Lripzig —Konigl. Sachs. Gesellschaft der Wissenschaften.
Litt“E.—Revue Biologique du Nord de la France.
Lonpon.—Royal Microscopical Society.
British Museum (Natural History Department).
MANcHESTER.—Microscopical Society.

Owens College.
MARSKILLE.—Station Zoologique d’Endoume.

Musée d’ Histoire Naturelle.

MassAcuusnrts.—Tufts College Library.
MECKLENBURG. —Vereins der Freunde der Naturgeschichte.
MELBOURNE.

Royal Society of Victoria.
MontTpreLiuirr.—Académie des Sciences et Lettres.
Mcscou.—Société Impériale des Naturalistes.
Nancy.—Sociéte des Sciences.

Napout.—Accademia delle Scienze Fisiche e Matematiche.
New Brunswick.—Natural History Society.

Oporro.— Annaes de Sciencias Naturaes.
Parits.—Museum d’Histoire Naturelle.

Société Zoologique de France.
Bulletin Scientifique de la Franee de la Belgique.
PHILADELPHIA.—Academy of Natural Sciences.
PitymMourH.—Marine Biological Association.
Sr. Louis, Miss. —Academy of Science.
Sr. Pererspure.—Académie Impériale des Sciences.
SANTIAGO. —Société Scientifiq du Chili.
STAVANGER.—Stavanger Museum.
SrockHoLtm.—Académie Royale des Sciences.
Sypnry.—Australian Museum.
Tox1o.—Inperial University.
Tor1no.—Musei de Zoologia ed Anatomia Comparata della R. Universita
Toronro.—Canadian Institute.
TRIESTE,

Societa Adriatica de Scienze Naturali,

LIBRARIAN’S REPORT. XXXll.

UpsaLta.—Upsala Universitiet.
W ASHINGTON.—Smithsonian Institution.

United States National Museum.

United States Commission of Fish and Fisheries.
We.iincTon, N.Z.—New Zealand Institute.
Wirn.—K. K. Naturhistorischen Hofmuseums.

K. K. Zoologisch—-Botanischen Gesellschaft.

ZurRicH.—Zurcher Naturforschende Gesellschaft.

:

sg | THE LIVERPOOL BIOLOGICAL SOCIETY.

Dr. In Account witu ISAAC C. THOMPSON, Hon. Treasurer. Cr.

1895. £
To Use of Rooms, &c., University College ............... 3
,, Lea and Attendance at Meetings........... EE Rae ies 4
», Printing Reports and Proceedings, T. Dobb & Co... 40
,, Binding Books, Jeans & Co............ St ae see oat ioat
Pe CCS MOTMIVe DOLCS sty in aac gee el siscata oitetoidisiu acenateane se 2
POSE CES® center wmiaat Miclaneideas «bcdse eleven Metiemcnic care vencan 4
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Pewalance im hand AUensh olst, VO9D o....:..ssaceree. 65

wo} ~~. oC ao A oS NN oS &
SOS oo Oo

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ISAAC C. THOMPSON,
Hon, TREASURER.

LIVERPOOL, August 31st, 1895.

1895. £ Ss...
By Balamee;-aucush alst 1894) snus. ensens eee Bananas ie
5, 8 Members’ Entrance Fees at 10/6 ............. Rete 4 4 0
,, / Student Members’ Entrance Fees at 2/6...... ne OFS 6
_, 58 Members’ Subscriptions at PAW a aseasccarnee tee 60 18 0
,, 21 Student Members’ Subscriptions at 5/- ............ Sao 10
£121 18 9

By Balancenin Unions bane sentence ent Sane tar £65 7 2

Audited and found correct,
ALFRED LEICESTER.

ee ee

. TRANSACTIONS
LIVERPOOL BIOLOGICAL SOCIETY.

INTRODUCTORY ADDRESS: “STRUCTURE
AN BUNCTION .”

By Francis Gotcu, M.A., F.R.S., President.

PROFESSOR OF PHYSIOLOGY IN UNIVERSITY COLLEGE, LIVERPOOL.

I HAVE first to express to my fellow-members of the
Liverpool Biological Society my sense of the honour which
the Society has accorded me in electing me President for
_ the session 1894-95, and my gratification at the emphasis
_ which this places upon those aspects of Biological study
and inquiry with which I am especially identified. The
erowth of science necessitates subdivision into departments
and tends in consequence to isolate the departments thus
made, so that Physiology, by a natural process of evolu-
tion, eradually diverges from those sciences which deal
- more especially with the structures of living things—
Zoology and structural Botany. The problems presented
to the physiologist, although fundamentally the same as
those offered by Zoology, are in their immediate aspect
different in kind; the methods employed for their elucida-
tion are, in many directions, special to physiological
workers and thus, even had there been no adventitious
circumstances to accentuate the divergence, a partial
separation would probably have occurred.

It is however hardly necessary for me to remind you

that adventitious circumstances have existed, and exist

still, which have effected, if not a divorce, at least a
judicial separation between the wedded studies of Physi-
1 ology and Biology. Pope crystallised, by the elegant
terseness of his diction, many a prevalent notion, and
amongst such is that expressed in the well known line,

2 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

‘““The proper study of mankind is man.” His contem-
poraries thought that such lucid phrasing justified them in
deeming him ‘‘the incomparable poet,” forgetful that
the aphorism expresses simply the prevailing idea of the
day, and that, as years pass, prevailing ideas change.
But Physiology has been and still is one aspect of Pope’s
dictum. It exists to-day essentially as Human Physiology,
the study of the normal functions of the healthy human
organs. It is true that this study is, in many cases,
carried out on the tissues and organs of a few lower
animals, but the facts thus revealed are used rather to
throw light upon the functions of our own organs than
to elucidate the functions of the animals themselves.
We are, and shall always remain, to ourselves the most
important of all living things, nor can we wonder at this
when we remember, that the whole world and all that it
contains has its existence, so far as we are concerned, in
our own enigmatical sentient being.

Physiology, the study of the functions of living things,
what they do, joined to a blind groping enquiry as to how
they do it, owes everything to Medicine. Its Laboratories
are adjuncts to Medical Schools, the subjects of mvesti-
gation in such laboratories are in the vast majority of
cases such as directly tend towards the advancement of
knowledge useful to Medicine. ‘The debt my science owes
to Medicine cannot be overestimated but like most debts
it leaves the debtor with the chains of his bondage still
upon him. In this case the links of this chain become
visible when we realise the extent to which physiology
has been divorced from that great science of which it
really forms one entire aspect—I mean Biology. I rejoice
therefore that though in more exalted associations,
Physiology has by the shere weight of its metal, been
compelled to constitute itself a different section to Biology,

OPENING ADDRESS. 3

yet, in this Society, the strength of the tie which must
always exist between the two aspects is acknowledged,
and is this year brought into prominence by the position
which [I fill to-night.

With these thoughts in my mind it seemed that I should
best discharge the first duties of my office by inviting
you to consider the close relations which must exist
between the two great aspects of Biological phenomena,
and I do so in the hope that both the physiologist and the
biologist may be led to the conclusion that a concurrent
study of these two aspects is essential if we would further
to our utmost the advance of biological knowledge. This
may seem as self-evident as an axlom, requiring no
argument to support it, no demonstration to enforce it;
but whatever the extent to which it is recognised by the
judgment, it is not adequately recognised in point of fact;
for where age the physiological laboratories for the system-
atic study of comparative physiology, side by side with
comparative anatomy? Isolated inquiries have been
undertaken, but systematic work and adequate equipment
for prosecuting that work on any one class of invertebrate
animals does not exist in connection with any Biological
or Physiological Department, not merely in this country
but in the whole of Europe and America, except perhaps
at Naples.

It is surely time to consider to what extent it is possible
to combine the simultaneous study of the function and
the structure, not merely of man and of lower animals in
relation to man, but of each group of living things as such.

In order to enforce this and show its value, I propose to
bring to your notice, some few typical instances, which
may serve to illustrate the enormous gain which a study,
such as I have indicated, would be both to the Biologist
and the Physiologist.

4 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

What does the examination of a structure tell us? It
tells us the mode in which the various parts of the animal
or plant are arranged, and the special features which
characterise their appearance. It tells us to what extent
construction resembles or differs in the various animals
and plants. It tells us the changes which occur in the
character and form of the whole or of special parts as these
develop from the first stage of their existence and grow.
From such records we infer the relationship of a structure
or an animal to its contemporaries and even to forms

‘which once existed in the past. The genius of Darwin

and his successors has awakened so intense an interest
in the working out of these relationships that recent
Biology seems to be swept away, as by an ocean tide, into
the sea of a vast speculative inquiry. Evolution is a
word to conjure with, and the part played in this by
Natural Selection, by Sexual Selection etc., the significance
or insignificance of acquired characteristics, whether or
no these can stamp the whole organism as with a die, so
that by heredity they are transmitted to a descendant,
such problems offer so fascinating and so extensive a field
for Biological speculation, that one side of Biological
inquiry, that, namely, which determines the structural
changes in consequence of growth either of the individual
or of the race, has become of late years predominant.
And yet such problems have been attacked almost entirely
from one aspect and no systematic attempt has been made
upon the functional side of the question—Would it not
therefore be wise if Biologists planned physiological
investigations which should, in the end, throw light on
the transmission or inheritance of functions as well as of
structures; but for this inquiry previous knowledge of the
functions is indispensible. In what respects do the func-
tions of the tissues of animals resemble or differ from one

OPENING ADDRESS. 5

another; that is the vast field which waits for systematic
investigation. Until that inquiry is made, ought we not
to regard the present evidence upon these intensely
interesting questions as incomplete? Yet a great deal
of Biological Study is devoted to the examination of
structure solely with the view of attempting to support or
repudiate speculative hypotheses which hover round the
great central truth of Evolution.

I do not propose to-night to refer again to this very wide
subject, both because of my incompetence to do so ade-
quately and because I wish to limit my range to subject
matter which involves more direct issues. I pass on
therefore to consider to what extent structure is a guide
to function.

First class of wmstances, inferring Function from
Structure.

Suppose that on dissecting one of the higher animals I
came upon a mass of tissue which in my opinion looked like
a muscle, and that on examining it carefully, not merely
with the naked eye, but also microscopically, 1 found
that it contained all the elements of striped or voluntary
muscle. Should I then be justified in stating what its
function was? Having ascertained the form does that
tell me the character of the living activity? I should
undoubtedly have very strong grounds for the inference
that this mass of muscle really was capable of contracting
in response to a stimulus, and thus moving the parts to
which it was attached, but I should not know it could do
so unless I saw it so act in the living state in response to
such a stimulus. Even having ascertained this, | am by
no means warranted in the next assumption that this
muscle which I have found to be supplied by nerves coming
from the central nervous system can respond, not merely

6 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

to the external stimulus I apply to it, but to the more
subtle and unknown stimulus which emerging from the
central nervous system courses down to it along its nerves.
The form is certainly there, but not necessarily the function.
Many of the muscles of our body will contract to a stimu-
lus and are supplied by nerves which reach them from
the central nervous system, but we cannot by the strongest
voluntary effort make them act. Indeed, even those
muscles which do so contract in response to the will act
but in a partial and incomplete manner, and more exact
inquiry seems to show, that although all the component
muscular fibres are structurally alike, they do not all
respond alike to the will stimulus—some remaining
quiescent. It would therefore be a much bolder assumption
to infer from the structure of a muscle the functional
characters of its activity in the living animal. Experi-
mental investigation alone can decide these.

Yet undoubtedly, on the discovery in an animal of a
mass of muscle, there is irresistibly forced on the mind
the conviction that since its structure and connections
resemble those of other muscles, its functional activities
must do so too. We jump from ascertained structure to
unascertained function, and why? Because a vast number
of instances of an accurate correlation of structure and
function force the judgment to this leap. What has been
so often must be again, and in the majority of cases will
be so again. There is thus a very great probability that
this jump is one on to sure ground, but it may be a
plunge into the abyss. This is the rock on which many
a purely structural investigator has suffered shipwreck.
A conviction has been forced upon his judgment such as
Wordsworth expressed in his Duddon sonnet. ‘The
Form remains the Function never dies.”’

Let us be perfectly clear that structure does not deter-

OPENING ADDRESS. 7

mine function; it limits it, it prescribes the mode in

which the activity shall especially manifest itself and that

_ only. Itis only in this sense true “that every appreciable
_ difference in structure corresponds to a difference of
function.’’*

The inference is, however, often correct and I will now
refer to those demonstrative examples of the correctness
of the inference and the dangers which beset it, which
are afforded by the examination of a class of special
_ organs—the socalled electrical organs present in certain
fishes.

More than a century agot it was discovered that the
peculiar numbing power possessed by certain sea fish,
(Torpedo or Electric Ray) was due to the power they
possessed of being able to discharge electric currents of
great intensity, these being generated in organs in the
body of the fish. These organs, when investigated by
John Hunter anatomically, were found to consist of piles
of thin plates arranged in columns and supplied by an
enormous number of nerves and nerve branches. When
a muscle contracts a great many things happen besides
the change in form and among these is the generation of
electrical effects; but one special aspect of its activity—
that of contraction—transcends all others. In the elec-
trical organ, an electromotive disturbance occurs in the
neighbourhood of each nerve ending and this now far
transcends any other aspect of its activity.

Another fish, this time a fresh-water one, the Gymnotus
or Hlectric Hel, found in the Orinoco, was known to
possess the power. When it was examined anatomically,
also by Hunter, it was found to have four enormous

* Sanderson. British Association, Address, Section D, 1889.
+See Phil. Trans., 1773: Discovery by Dr. Walsh, p. 461, and Anatomical
investigations by John Hunter.

8 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

organs occupying almost the whole of its body, each
composed of similar piles of thin plates arranged in
columns. Still another fish was known to have the power,
the Malapterurus found in the rivers of Africa; when
examined anatomically its thick skin was seen to consist
of piles of plates, each pervaded with innumerable sub-
dividing nerve-fibres. Organs thus built up and possessing
these properties were termed from their function Electric
Organs.

Now it was felt by Mr. Darwin (and indeed is still felt
by all Biologists) that the rare possession by fish of organs
so complicated yet so effective, was a formidable fact which
the theory of Natural Selection was unable adequately to
erapple with. In what way could such organs have come
to be in the possession of these special fish and their
descendants? Surely their near relations might be reason-
ably expected to have them also.

The common Skate is such a near relation of the Torpedo
from the point of view of descent. Mr. Darwin therefore
suggested that it should be examined for the purpose of
ascertaining whether such an organ existed init. In 1861
McDonnell of Dublin* announced the discovery of such
an organ. Upon what grounds? These two only; first,
that in the same anatomical situation as in the case of the
Torpedo, there was a mass of substance containing columns
and an extensive nerve supply, and secondly, that on
histological examination this structure had a large number
of separated tubes with a large number of nerve endings. |
He jumped at the conclusion that the organ was an
electrical one and Mr. Darwint+ accepted that conclusion.
But they had jumped into the abyss. The organ was not
an electrical one. The same mistake had been made in ~

* McDonnell. Electric organs of the Skate. Nat. Hist. Review, p. 59.
+ Darwin. Origin of Species, (p. 150, 6th edition).

OPENING ADDRESS. 9

a 1801 by St. Hilaire and repeated by Meyer in 1843 and

de Lamballe in 1858. In 1822 Blainville pointed out the

fallacious character of the inference. The organ had been
correctly described some years before and Leydig in 1850
showed that it forms a central terminus for a series of
glandular tubes which run over the surface of the body of
the fish. It is incapable of generating any electromotive
change in response to a stimulus, and McDonnell’s infer-
ence was therefore unsound. Here we have an instance
of the failure of the inference from supposed structure
inadequately examined. The organs in question form
part of what Leydig and others have called the sensory
canal system. You see the difficulty of getting clear of
this tendency to infer function. It 2s a canal system, but
as far as I can make out the only evidence in favour of it
being a sensory canal system is that the nerve which supplies
it, is said to be the 5th cranial which, in higher animals, is
mainly a sensory nerve, and that cells resembling those in
sensory structures are present in some of its terminations;
but there is no physiological ground that I know of for the
functional assumption. The activities of the organ and of
the gland tubes connected with it are to this day unknown,
since appropriate physiological experiments have not yet
_ been devised to ascertain them. Yet unwarned by the
experience of de Lamballe and McDonnell the structure
has been very generally given the functional attributes of
_ aterminal sensory organ. It may be merely secretory,
and indeed Fritsch and others have urged this, but let us
not commit another jump into the abyss; physiological
enquiry in conjunction with anatomical can alone deter-

mine what these organs really do.* (See Fritsch; Berlin

*In the Section of Biology at the meeting of the British Association,
Oxford, 1894, Mr. Collinge brought forward anatomical evidence to show
_ that the nerve supplying the canal system of Elasmobranchs is not the 5th
but the 7th Cranial Nerve, i.e., an efferent or motor one! !

10 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Acad., 1888, VII, page 301, and Ewart: Transactions of
Royal Society, Edin., 1892).

Seventeen years before McDonnell’s mistake Dr. Stark,
of Edinburgh, discovered in the same fish (Raza batis),; but
in quite a different situation (in the long tail of the Ray)
an organ with piles of plates arranged in columns and
with a very extensive nerve supply. He inferred from its
structure that it was an electrical organ, but was careful
to say that he had no evidence except its anatomical
resemblance to the powerful active organs of the Torpedo.*
This organ was examined later as to its function and was
found to be capable of generating electrical currents of
considerable intensity, though not strong enough to be
perceptible to a man’s hand, except perhaps in the case of
very large fish. Stark’s inference was sound, because in
this case the function, when investigated, was found to be
that which the structure led its discoverer to predict.

Since then a number of fish in the Nile, one especially
the Mormyrus, have been found to have similarly con-
structed organs, but for a long time these were termed
pseudo-electric organs. They have now been examined
by physiological methods, and undoubted evidence of
electrical changes of considerable intensity has been
obtained. They are therefore true electrical organs.

It is worthy of note that in both this case and the skate,
we should never have got at the function but for the
discovery of the structure. These are instances of like
function being inferred from like structure and they
illustrate at once the great value of the structural resem-
blance in guiding us to a true functional interpretation,
and the dangers which the inference involves when
unsupported by physiological inquiry.

* Stark. Proceedings Royal Soc., Edinburgh, Vol. II, No. 25. This
discovery appears to have been disregarded for many years.

~

OPENING ADDRESS. Att

_ There are heaps of such things waiting for investigation.
Such are the so-called special sense organs of invertebrates;
_ the so-called ‘‘ pineal eye”’ of certain lizards lying in the
- middle line and now concealed by the epidermis. This
last is a structure with a lens and a retina-like arrangement
of cells; its whole formation, its connection with the
nervous system and the corroboration furnished by
palaeontological inquiry have led irresistibly to the con-
clusion that it is an eye. Yet the conclusion is an
_ inference unsupported by Physiological experiment, and
| though probable justifiable, not necessarily so. Its
| function has never been ascertained, and, as far as its
_ structure goes, it might possibly be a heat sensory organ
~ not a light one.

like structure, therefore, leads us to postulate like
function but the character of the function is an inference
to be verified or not by physiological methods, and by
_ these only.

Second class of instances, inferring changing Function
from changing Structure.

_ So far I have only referred to a definite structure and
the definite function associated with it. The whole
- question of the correlation becomes more complicated
- when we have to deal with a structure, which in the
course of its development undergoes a gradual meta-
-morphosis, the mference being that the function undergoes
a similar change.

Here again the inference may be justified by verification,
but without ascertaining the function it remains an
“assumption, and a much greater one than was the case in
those instances just referred to.

To illustrate this I might select almost any organ in
different stages of its development, but a very beautiful

12 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

example is afforded by the electrical organ to which I
have already referred. The stages of growth in the
developing Skate are so slow in time, that its tissues are
elinently suitable for such enquiries.

To take the case of the electrical organ, biologists have
endeavoured to answer this question—through what stages
of form did the tissue pass before the present electrical
organ was evolved ?

The answer to this, in the case of the Skate, is obtained
by the study, first, of the electrical organs in different
species of Skate, secondly, of the electrical organs in one
species at different stages of its own growth. Both
investigations reveal the fact that as far as form is
concerned, the early condition is that of a series of muscle
fibres, and that these dwindle down to form the plate
whilst the nerves to the muscle remain, and by sending
out branches in repeated subdivision form the dense
nervous net work.* Species are found in which the

different plates of the organ present a curious transitional __

condition between a muscle fibre and an electrical organ

plate. Such are the plates of the Raia radiata; histological

specimens show that the structure is still there but quite
modified in form. Recent microscopic investigations by
Engelmann under polarised light seem to show that such
parts of the muscle as act in a special way on the
light (doubly refracting), these parts occurring in bands |
along the muscle fibre, gradually decrease in amount,
whilst the adjacent singly refracting substance remains. t
With all this, what is the change in function? It remains
at present unknown; all that I know is that in such an
imperfect organ, composed of half muscle and_ half
* Ewart. The electric organ of Raia radiata. Phil. Trans., Vol. CLXXIX,

p. 539. .
+ Engelmann. Pfliigers archiv., Vol. LVII, p. 149. (1894).

OPENING ADDRESS. 13

electrical organ, the electrical effects were, in the single
instance which I have examined, extremely small, far
smaller than those which normally accompany the activity
of even the sartorius muscle of the frog, and that on the
other hand, this transitional structure, whilst it had no

electrical function to speak of, had in this particular case

no contracting function that could be detected by our
present methods.

If this is so, it is obviously an unsound inference to
assume that this structure is one in which the function of
the two forms, those of muscle and electrical organ, are
represented ; they both appear to be non-existent.

Here then alteration of structure has enormously
modified the function, but apparently in such a way as to
abolish its former predominant characteristic without
introducing that new characteristic which the newly
developing form would lead one to infer as being present.

I bring forward this instance because of the essential
importance to any inquiry as to the structural changes in
living tissues due to their growth, of simultaneous
investigation into the alteration of the functional activity.
Comparative physiology of this kind would thus be of the
very greatest value for the correct interpretation of the
phenomena of growth and for the determination of the
question, as to how far it is the change in structure which
prescribes the change in function. Perhaps the change
in function may be the exciting cause for the alteration in
structure, for the force which we term “ vital”? sways the
mechanism which is after all nothing but it’s tool.

Third class of instances; inferring excitatory
Functions from Structures.

If structure is not a sure guide to function even when
this function is definite and comparatively simple, it must

14 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

be a far more unstable guide when the function is both
more general and more varied. This last is the case when
we have to deal with a plurality of functions, any one of
which may be in the ascendant in a particular part of the
tissue without any clear structural reason to hang that
ascendancy on.

Let me remind you of one typical example. In the
Dionzea muscipulata, the two lobes of the leaf are
sensitive to mechanical contact and other stimuli. When ~
touched, they rapidly close and imprison, as in a dungeon
with barred gates, any object which is in contact with
their inner surfaces. This movement is produced when
the leaf is rudely touched, but there is one part which is
exquisitely sensitive to the lightest touch; this part is in
the middle of the leaf in the immediate neighbourhood of
three hinged hairs, and is termed the trigone.

A fly walking over the leaf does not excite the closure
of the lobes until it reaches the central trigone, or strikes
these so-called sensitive hairs. The use of this to the
plant is plain on the assumption that it is desirable for it
to have an occassional meal of animal food; for the fly is
so far in that it cannot get out before the gates close, and
implanted in its prison it is slowly digested by the peptic
juices which are poured out upon it by little glands on the
inner surface of each leafy wall.

Surely with such an apparently elaborate function one ©
might expect that the structure of the cells in this sensitive
part would be different to the structure elsewhere.
Histological investigation, however, shows no difference
corresponding with the ascendancy of the function. The
sensitiveness (or excitability) is but one of a plurality of |
functions possessed by the cells (power of retaining liquids;
loss of this power on stimulation, electrical changes, etc..,)
and is the common property of all the cells in the leaf, but

OPENING ADDRESS. 15

in this region, at the base of the hair, it is enormously
in the ascendant. Structure is no guide at all to this
ascendancy, because the former remaining apparently the
same, the latter alters. This 1s the case with most of the
so-called excitatory phenomena. The whole of physiology
simply bristles with instances; thus, a muscle is acted
upon by a certain salt—sulphate of veratria ;—its micro-
scopic structure is apparently unaltered, but its activity is
changed so enormously, that when aroused by a single
stimulus, instead of shortening and resuming its length
again In yy, 1t now contracts and remains unable to relax
for almost 1 minute. |

Again, a nerve, if exposed to weak ether vapour, suffers
no perceptible alteration in structure; but as long as the
vapour is present it ceases to carry out that ascendant
phase of its activity for which it 1s really framed; it cannot
pass on the mysterious awakening which, flashing down
through each successive part, is called a nervous impulse.
On removing the ether, back comes this conducting
property so that the vapour has not permanently injured
the nerve. All the phenomena of anesthesia are exten-
sions of such a case as this.

I might multiply examples almost indefinitely; I should
simply be stating the functions of the excitable tissues and
the circumstances which modify these, for in very few
instances are such modifying circumstances sufficiently
gross to cause any perceptible alteration in structure part
passu with the profound alteration they produce in
function.

Fourth class of instances ; inferring Structure from
Function.

This is an inference of precisely opposite character to
those hitherto considered and is eminently the property of

16 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Physiologists. Although a most fruitful one it is certainly
dangerous and it has often happened that the Physiologist,
relying upon it, has leaped into the abyss.

I need hardly remind you of the innumerable instances
in which a structure has been inferred from the observation
of a function. Thus, a tissue is ascertained to swiftly
alter its form in response to a stimulus and we infer that
in this part will be found muscular fibres. The judgment
is now urged to deduce the structure from the function.
In a vast number of cases the inferred structure is
subsequently found to be present, but it may not be found,
and then the only safe course is to acknowledge that an
inference has been made and to wait. In Biology, more
than in any other Science, we must possess our souls
in patience.

One case, in which such an inference wrongly made has
bothered the Physiology of one particular organ for years
is that of the iris. The iris of the eye is an organ with
an aperture, the pupil, which increases and diminishes in
size. It decreases when light falls on the eye, when we
look at a near object etc. Its size is constantly swaying »
in response to nerve impulses which reach it. By what
mechanism is this movement effected? The inference is
that itis done by muscle. Anatomical examination shews
that the iris contains fibres disposed in circular rings
round the pupil; these are seen by microscopic examina-
tion to be muscular fibres; the contraction of these
will therefore diminish the size of the pupil, and when
the contraction decreases the pupil will expand. Now, no
other muscular fibres were definitely found in the iris of
mammals, so the dilatation of the pupil was considered to
be in all cases due to the cessation of the contraction in the ©
muscular ring or sphincter pupillae, and the iris was said
to expand passively. There were however many physio-

OPENING ADDRESS. 17

logical facts which seemed to indicate, that the pupil
might be made to expand even when this sphincter was
not affected. Some physiologists therefore demanded a
second radial muscle, which by its contraction could pull
out the pupil—a so-called dilator muscle with radial fibres
like spokes in a wheel. Although such fibres exist in the

birds eye, none were found in the mammalian iris, and it
was therefore thought to be vain to assert from experiment
their presence. The observed phenomena must, it was
said, be capable of some other interpretation, active
expansion was thought to be impossible. In 1892, a
discovery made simultaneously and independently by Lang-
ley* and Heese+ showed definitely that in the cat the
iris could be made to expand actively. If its edge be
stimulated by suitable electric currents led through the
corneo-sclerotic junction the pupilis not constricted but
expands at that point and by shifting the site of the stim-
ulation this local expansion is similarly shifted. Careful
examination of the iris shows that two movements occur
in it, each confined to the neighbourhood of the stimulated
part. Immediately the stimulation begins there occurs a
localised puckering of the pupil all radial lines at this
point being brought visibly nearer one another; this must
be due to the contraction of the circular sphincter. As
the stimulus is kept on, there is suddenly seen a new
puckering causing folds which are piled up near the edge
and are circumferential, whilst the pupil at this point
expands. The two movements are now seen to be going on
in the iris at the same time. After the stimulation has
ceased, the sphincter puckering, which was the first to begin
stops, and now the second series of folds is seen by itself.
This is a demonstration ad oculos and in oculo, that there

* Langley and Anderson. Journal of Physiology, Vol. XIII, p. 555.
+ Heese. Pfliigers Archiv. Vol. LII, p. 535.

18 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

is in the iris some radially disposed structure which by its
contraction dilates the pupil and causes this second set of
folds. Why then did the older physiologists give it up and
even consider themselves wrong? Because they had
deduced a structure from the function and assumed that
this radial structure must be a conspicuous muscle. There
is no conspicuous radial muscle, but a more careful exam-
ination by Heese has shown isolated radial fibres, though
whether muscular or not we do not yet know.

Function here has guided finally after many failures to
structure. Mistakes may occur, but the advantage of
inferring the structure from the function is often quite as
great as the reverse inference. 'T'o it we owe the dis-
covery of structural changes underlying the chemical
processes of living cells.

Fifth class of stances; changes in Structure discovered
to underlie changes in Chemical Actwity.

For years microscopic investigation in conjunction with
Anatomy was the sole guide in the endeavour to find such
changes, but a time came when this investigation unaided
by a knowledge of the function seemed to be insufficient.
To make this clear, take the activity of secreting cells.

The salivary glands, pancreas, stomach, liver, etc., each
secretes its own special digestive fluid. The juices, and the
antecedents of their constituents, are evidently manu-
factured in the cells of each special gland, and such cells,
when examined microscopically, showed characteristics
which distinguished them from others. What, however,
are the structural features in the cell which are associated
specifically with the manufacture? ‘To this question
the older Histology could give no satisfactory reply; it
seemed almost beyond the limits of histological manipu-
lative possibility to even give us alead. Then the subject

OPENING ADDRESS. ~ 19

was approached from the functional side. Suppose the
gland and its cells are worked so hard that they are
exhausted of their potential secretory activity, and they
are then compared as to structure with others in the full
vigour of secretion. Surely they must look different,
especially if they are examined in the living state.
The pancreas of the rabbit was so examined by Kuhne
and Lea, and it was observed that the exhausted cell
was small, whilst the manufacturing cell was not only
large but full of granules.

If the cells are fixed by microscopic reagents, then, as
a rule, they are altered; the older methods of hardening
tissues reduce their cells all more or less to the same
common denomination. By the use of osmic acid, alone
‘or in combination with various reagents, it 1s now found
possible to fix them so that their death will not essentially
alter their living appearance, and structural features have
now been found in the pancreas, the salivary glands, the
liver, the gastric mucous membrane, etc., which features
vary part passu with the variation in the chemical pro-
perties of their secretion. Instead of Physiology following
in the wake of Histology, it has now become the guide,
pointing the direction which microscopic work should take.
As Sanderson said in his Newcastle address to the
Biological Section, at the British Association, in 1889:
“Whereas hitherto the greater part of the work has
consisted in the interpretation of facts arrived at in the
first instance by anatomical methods of research, Histology
once the guide of Physiology has now become her hand-
maid.”

I can think of no recent instance more striking than
that of the structural changes in the cells of the mammary
gland accompanying the various phases of the secretion
of milk. Milk contains all the essential constituents of

20 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,

animal food, proteid, albuminoid, fat, sugar and salts, and
they are elaborated in these cells. Such elaboration
should have at the back of it, definite structural changes
in the cells. It seems clear from recent research that,
when the secretion is being formed in the gland, the
mammary cells alter not only by duplication of their
nuclei but by the appearance of specialised chromatin
filaments in the cell-substance. Simultaneously with
these, fat globules appear in juxtaposition with the new
nuclei and with these filaments, and finally the new
nuclei, the filaments and the fat are all discharged from
the cells into the duct. Here the nuclei and filamentous
particles appear to pass into solution, the fat remaining
as a fine emulsion.*

Sixth class of instances ; structural changes associated
with nervous actiwity. |

In nervous tissues, by following the same inference from
function to structure, definite alterations have now been
demonstrated. Let me confine myself to one class of
nervous tissues, that associated with vision.

The nerve-fibres from the retina of the eye start from
cells there and stretch back towards the brain, where they
end in the immediate neighbourhood of other cells, from
which other nerve processes spring. Through all these
the mysterious change which we call a nervous impulse is
conducted, this in the case of the eye being aroused by the
action of light.

Physiological experiments have shown that such nerve
impulses pass through or into special masses of cerebral
nerve-cells, localised at the base of the brain in the corpora
quadrigemina, etc., and on the surface in the occipital

~*See Steinhaus Die Morphologie der Milchabsonderung, Du Bois Archiv
f, Physiol., 1892, Supplement, p, 54. Jeti

OPENING ADDRESS. 1

lobes of the cerebral hemisphere. Histology, unaided, could
find no change in these cells in consequence of nerve
impulses having followed one another through this nervous
chain, but lately structural alterations in these cells have
been ascertained, the road for this discovery having been
made by the more accurate physiological knowledge of
cerebral function which physiological experiment has
revealed. If one eye of an animal be bandaged, the
animal exposed to light and then killed, on examining the
nerve cells in the retina, and in the particular regions of
the brain just referred to, it is seen that the cells on the
side of the bandaged eye are quite different in appearance
to those on the other side. ©

Those through which the nervous impulses passed show
alterations both in the size of their nuclei and in the
micro-chemical characteristics of their protoplasm. This
research, hardly yet completed by Dr. Mann of Edinburgh,
is undoubtedly the initial step of a great histological
inquiry, and to this field physiology has guided the micro-
scopist ; function has thrown light upon structure.*

That there is an inference in the above, is no doubt true,
and this is well illustrated by the last instance to which I
can allude to-night—one of the most brilliant of all the
discoveries connecting function and structure.

The sense organs themselves, not merely the nerves and
nerve-cells, but the cellular apparatus which is acted upon
by the external force, which, in the case of the eye,
transmutes in its living crucible waves of light into visual
nerve impulses, this peripheral organ changes in structure
in consequence of the action upon it of the external agent.

If the frog’s eye be exposed to light, and the retina
contrasted as to structure with an eye kept in the dark,

* Mann, Changes in Nerve Cells, Journal of Anatomy and Physiology,
Vol. XXIX, p. 100.

22 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the cones and the pigment which form the receiving
surface for the light are found to be different in the two
cases.* Hngelmann discovered, that the stalks of the
cones exposed to light shorten, and that the pigment
cell processes stream down deeper into the retina. The
distinguished physiologist, who ascertained these changes,
guarded himself against the fallacy of a too sweeping
inference. He ascertained whether it was necessary that
the light should reach the retina to produce the effect.
For this purpose, the liead of the frog was enclosed in an
absolutely light-tight bag, and light was allowed to shine
on the skin of the body. The wisdom of such an experi-
ment was justified by the result. A similar definite
change, but less in amount, was found, although the light
could not enter the eye, and could not therefore act
directly upon the retinal epithelium.

Now the skin is well known to be sensitive to light, as
is shown by the changes this produces in its pigmentation.
The discovery, therefore involves more than appears at
first sight, but I need hardly say that it is, on that account,
still more important.

CONCLUDING REMARKS.

All these instances show us how pointless is structure
apart from function, and how baseless and unstable is
function apart from structure, and make in my judgment
an irresistable argument for the simultaneous examination
of both.

Why is not such an examination more widely carried out
in Zoology ? Is it not because of that judicial separation
between comparative anatomy and physiology, which, as
was pointed out early in this address, is largely due to

_ * Engelmann. Pfliigers Archiv., Vol. XXXYV, p. 453,
Nahmmacher. Physiol. Labor, Utrecht (1898).

OPENING ADDRESS. ay OS

Medicine? That this is one cause seems to be probable
_ when we look at the state of botanical science. Physiolo-
gists do not as a rule meddle with vegetable physiology,
and I remember when I had the honour of assisting my
teacher, Professor Burdon Sanderson, in his experiments
upon the excitability of the Dionza plant, being asked
““why we were working at vegetables and what they had
to do with Human Physiology.”’

The Botanist has to work out his own physiological
salvation in fear and trembling, but it is undoubtedly for
the good of that science, since the study of structure side
by side with function is the true method. As a result
botanical science has its vegetable Physiology, but where
is the Physiology of the Invertebrate ?

- Comparative Anatomy and Animal Physiology must
both suffer if they allow the separation between them to
become a divorce; they should be wedded, that, from their
union may spring as their first-born, a new department,
devoted to the systematic study of Comparative Physiology.
What does the establishment of such a department
demand? ‘The needs are those which have to be met in
connection with all physiological work, but associated now
with such special zoological requirements as are necessi-
tated by the special direction of this inquiry.
Remembering that the function of the structures of the
lower forms of animal life, is the goal towards which all
work in this department must press, the question admits
_ of some such answer as this. There are three practical
methods for the study of the physical aspects of living
things. These are either chemical, mechanical or
microscopical. A comparative Physiological Laboratory
would need adequate equipment for chemical investigation,
for ascertaining such physical changes as movement,
electrical effects, heat effects and the like, and for exam-

94 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

ining the microscopic features presented by living, as
compared with dead tissues.

Comparative Physiology would however be nothing —
unless it were experimental, and for this the chief require-
ment is living material for experiment. Hence the
department needs such zoological equipment as will enable
it to keep living animals. Since a vast number of lower
animals are oceanic, this second requirement is to some
extent fulfilled in marine biological stations and all that is
necessary is for such stations to awake and realise the full
importance of this ideal Laboratory. They would then
be fitted with well equipped Physiological rooms as adjuncts
to their own buildings. Attempts in this direction have
been made with more or less success at Naples, and a
spasmodic effort was made by Paul Bert at Arcachon in
France, but in this last case, though the attempt yielded
an initial brilliant discovery in connection with the
respiration of fishes, it has since languished.

There are two difficulties—first and foremost, the most
important of all agencies has to appear. The new man.
It is not easy to find the comparative Physiologist, who
has been highly trained in both zoological and physiolog-
ical methods and knowledge. But I do not despair of his
advent; in the universities to-day are some who pass from
advanced zoological study, through the most extensive
physiological course which these universities can offer,
and I anticipate that in the future zoologists will be more
and more anxious to perfect themselves in the knowledge
of the methods used for investigating thefunctions of living
things. For I am certain that an immense tract of unex-
plored country with all its fascinating unknown lies open
to him who thus armed enters upon the quest. The second
obstacle, and that a serious one in this commercial country,
is the truly British stumbling block of expense. The

OPENING ADDRESS. 25

subject is not one which offers to the layman any definite
commercial advantage, so that, as far as England and
_ America go, it needs for its support that pious founder
whose arising even a Bodleian Librarian has now anxiously
} to await. Butas Biologists we need no such meretricious
stimulant as pecuniary gain; it is enough for us to know
that we shall obtain by Comparative Physiology a deeper
and truer insight into the complicated skein of the organic
world; we have our reward in the endeavour which we
make, either to trace or to realise that harmony which
~ underlies all the varied tints of a life, that
‘Like a dome of many coloured glass
Stains the white radiance of Eternity.”

26

EIGHTH ANNUAL REPORT of the LIVERPOOL
MARINE BIOLOGY COMMITTEE and their
BIOLOGICAL STATION at PORT BRIN.

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

DERBY PROFESSOR OF NATURAL HISTORY IN UNIVERSITY COLLEGE, LIVERPOOL 5
CHAIRMAN OF THE LIVERPOOL MARINE BIOLOGY COMMITTEE,
AND DIRECTOR OF THE PORT ERIN STATION.

[Read 9th November, 1894.]

THE work at Port Erin continues to flourish, and the
investigation of the Irish Sea generally is increasing in
scope. The laboratory has been well used during vacation
times; there have been a considerable number of dredging
expeditions, and several new lines of investigation have
been started which promise well. As there is much to
report upon, I shall pass at once to the usual statistical
statements without further introduction; and for informa-
tion in regard to the foundation and equipment of the
Biological Station, and as to the aims and methods of the
Committee, reference must be made to former reports.*
As usual, I am indebted to several of my colleagues on |
the Committee and to workers at the Station for kind
help which they have given me in the preparation of this
annual record.

STATION RECORD.

The following naturalists have worked at the Station
during the past year :—

DATE. NAME. WORK.
March. I. C. Thompson... ts ane eae Copepoda.
— W. A. Herdman oe Peres eee Tunicata.

_ C. Aa Einarst os eee Mes 559 on Collecting.

* Copies of the Sixth Report (56 pp. and 6 plates), in stiff boards, and
containing an account of the opening of the Station at Port Erin in 1892 by
Governor Walpole, can still be had, price one shilling each, on apphen to
the Hon. Treasurer.

W. I. Beaumont
F. W. Gamble ...

_.E. T. Browne
_ Alfred Leicester ...

F. E. Weiss
i Eiek »,...

F. E. Weiss
E. T. Browne
W. 1. Beaumont

E. T. Browne

H. C. Chadwick...
W. A. Herdman
I. C. Thompson ...

T. S. Lea. ;
Miss L. R. Thornely
Miss R. Alcock ...

I. C. Thompson...

A. O. Walker

W. A. Herdman

W. E. Ritter, California
A. Leicester

R. Hanitsch

W. A. Herdman...
I. C. Thompson...
Alf. Leicester

T. S. Lea

Arnold T. Watson
P. M. C. Kermode
G. W. Wood

. F. Gilchrist
. J. Halls

. A. Herdman

anitsch

. F. Gilchrist

Sea Sai
Foy

R. J. Harvey Gibson
Arnold T. Watson

MARINE BIOLOGICAL STATION AT PORT ERIN. 27

Nemertida.
Turbellaria.
Meduse.
Mollusca.
Algee,
Algee.

Algee.
Medusz.
Nemertida.

Meduse.
Synapta, &c.
Tunicata.
Copepoda.
Photographing Algz.
Polyzoa.
Tunicata.
Copepoda.
Amphipoda.
Tunicata.
Tunicata.
Mollusca.

Sponges.

Tunicata.

Copepoda,

Mollusca.
Photographing Alg.
Annelids.

General.

General,
Opisthobranchiata.
General,

General.

Sponges,
Opisthobranchiata.
Generale
Copepoda.

General.

Ceneral.

Algee.

Annelids

28 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Although this list is a somewhat shorter one than that
in last year’s record, still several of the workers stayed for
long periods, and so the amount of work carried on in the
laboratory this year has probably been greater than in any
previous year. Some of this work will be referred to later
on in the report.

Besides these workers, there were many visitors both
to the Laboratory and also to the Aquarium, which seems
increasing in popularity. During the season several
demonstrations on the animals living in the tanks were
given by the Director, and in September a special meet-
ing of the Isle of Man Natural History Society was held
at the Biological Station, when the Director gave an

_address on “‘ The Exploration of Our Coasts.”’

THE AQUARIUM.
A full account of the arrangement of the tanks and
pipes in the Aquarium-house was given in the last report.
Several minor additions and improvements have been

- made during this year. High stands for small aquaria

and glass vessels have been erected in front of the
windows. ‘The long table has been fitted with a leaden
covering, with upturned edges and waste pipe, so that
water spilled or leaking from aquaria may run off without
wetting the wood. New wall cases and shelving have
been added in the upper room for the reception of the
collection of Manks Marine Invertebrates (Echinoderms,
Zoophytes, Crustacea, and Mollusca) kindly presented by
Mr. G. W. Wood, and other similar dried or “ spirit ”
specimens collected in the neighbourhood. In this way
the walls of the room are becoming covered with collec-
tions which form an interesting little museum of local
marine zoology, and are a useful addition to the living

animals in the tanks when explaining the results of our
dredging investigations to visitors.

MARINE BIOLOGICAL STATION AT PORT ERIN. 29

The water motor and pump are not as satisfactory as
could be wished, but with a few alterations, which are
now contemplated, and which can be easily carried out,
we have reason to believe that the apparatus will work
better. Even without the pump we have, however, the
sea-water brought fresh every tide into our building by
means of the pipe from the beach and our deep well,
so that it is no difficult matter to keep the tanks supplied.

The Aquarium was in excellent condition during August
and part of September, under the care of Dr. Hanitsch,
and was open daily to visitors. The greatest number of
visitors in one day was twenty-nine. With a permanent
Curator, who will have fixed hours for going round with
visitors and demonstrating the contents of the tanks, this
department of the Biological Station will probably undergo
considerable development, may form a not inconsiderable
source of revenue to the institution, and will prove an
additional resource and attraction to the visitors to Port
Erin.

Amongst the more interesting or rarer animals which
lived in the Aquarium during the summer were :—Synapta
imherens, Porania pulvillus, Asterina gibbosa, Brissopsis
lyrifera, Panthalis oerstedi, Calocaris macandee (several
individuals of this deep-sea prawn lived for some weeks ;
they were partly covered with fine colonies of the rare
Polyzoon Triticella boecku), Hbalia tuberosa, Sarcodictyon
catenata (both red and yellow varieties), Alcyoniwm digi-
tatum, Scalpellum vulgare.

One specimen of Aurelia aurita lived in a tank for
over five weeks, but diminished steadily in size during the
time. At the end it was about half of its original diameter,
but still quite active and apparently healthy. The tank
given up to Shrimps, Prawns, and Mysids proved a success,
and was a great source of interest to visitors on account

30 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of the lively scenes that took place when the animals were
fed. There were always a number of the ordinary shore
animals, usually some beautiful vessels of brilliant ane-
mones from the Calf Sound, sometimes jars of Copepoda
and other surface organisms, while Hermit crabs,the sea-
mouse, a few flat fish, brittle stars, and sea-urchins were
always interesting to watch, and could readily be made to
lend themselves to purposes of instruction.

Besides its distinct use to the general public, our
Aquarium has another function. Several of the naturalists
at work this year have made more or less use of it, and
it has been felt to be a distinct aid in making several
interesting original observations, such as those on Pan-
thalis described further on. A number of animals
spawned during spring or summer in the tanks, and an
Actuma mesembryanthemum produced in March about 50
young anemones, which scattered themselves over the
tank and throve well.

TEMPERATURE OF THE SRA.

As we have had no permanent resident Curator during
the year, the sea temperatures have not been taken with
regularity, but several of the naturalists working, who
have been interested in the matter, and notably Mr.
HK. T. Browne in the earlier part of the summer, and
Dr. Hanitsch, who acted as temporary Curator during
the later part of the season, have kept daily records during
the period of their stay. These are entered in the
diary at the Station, and they show that, as in the case
of last year’s series, but not quite so markedly, the
temperatures of the sea-water in gradually rising as the
spring and summer advance lag slightly behind the in-
increasing temperatures of the air. This summer the
highest temperature (62° F., on August 21st) was ten _

ee ee *
; ;

_ MARINE BIOLOGICAL STATION AT PORT ERIN. ol

degrees lower than the highest of last year (72° F., on
August 15th), but the latter is probably to be regarded as
an exceptional occurrence, in an exceptionally hot sum-
mer. It is interesting to notice that in a few of the
observations taken by Mr. Browne in April, the water
out in the Bay, at a depth of 3 fathoms, was about 5° F.
below the surface temperature at the same time (after-
noon).

Nothing has yet been done in the direction of fish
hatching and lobster culture, for both of which important
industries Port Erin is so eminently suitable. There
are unfortunate parliamentary restrictions which at pre-
sent prevent the Lancashire Sea- Fisheries Committee
from spending money in this direction, upon what, from

their point of view, is foreign soil. The Manks Legis-

lature is, however, now stirring in the matter of their
coast fisheries. A bill has been promoted, has passed
through the House of Keys, and was lately promul-

gated, conferring powers, to make fishery bye-laws and

other necessary regulations, upon acommittee. It remains

to be seen whether this committee will content itself with

“rvestrictive’’ legislation, or will supplement that by the
at least equally important and necessary ‘“‘ productive ”’
work which leads to the encouragement and advance of
fish culture and the stocking of beds and banks. Such
direct action in aid of the fishing industries cannot be
much longer delayed, and I am now of opinion that our
Station at Port Erin should, during the coming season,
be utilised for experimental work on fish hatching. Hven
if we cannot obtain any assistance from the Lancashire
Committee, and if the Insular Government find they are
unable to aid in fish production, still our L. M. B.
Committee might establish one or two hatching boxes in
spring, test the quality of the water by an actual experi-

32 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

ment in hatching, and show what can be done even on a
very small scale and with very limited resources. |

DREDGING EXPEDITIONS.

During the year 1894 the following dredying expeditions
in steamers have been arranged, partly with the help of
the small Committee of the British Association alluded to
.n former reports. This B. A. Committee reported to
the Oxford meeting of the Association, and was then —
re-appointed for another year, with the addition of Pro-
fessor Weiss to cover the Marine Flora, and of Mr.
Clement Reid to aid in working out the Geology of the
floor of the sea :— }

I. March 20-25th. At Kaster the Committee spent some
days in shore-collecting at the southern end of the Isle of
Man, and hired the steam trawler ‘“‘ Lady Loch” for
two days’ dredging. On the first day the floor of the sea
to the north of Port Erin from Fleshwick to Contrary
Head at Peel was worked at twelve stations within four
miles of the coast, and at depths from 10 to 20 fathoms.
On the second day nine stations off the west of the Calf
Island at depths from 20 to 25 fathoms were dredged.

March 24th. 1. West of Fleshwick Bay, a quarter mile
off shore, 18 fathoms; bottom fine sand and broken
shells, with* Cliona celata, Gemellaria loricata, Canda
reptans, Ophwura ciliaris, Galathea intermedia, Portunus
arcuatus, Aporrhais pes-pelicam, Trochus magus, Ascidia
virgined..

9. West of Fleshwick, further north, half a mile off

* The few species picked out for mention in each haul are not to be
regarded as the rarest forms observed. In some cases they are the com-
monest. They are the forms which at the time seemed to us the most
conspicuous and characteristic of the haul—the most noteworthy inhabitants
of the ground,

MARINE BIOLOGICAL STATION AT PORT ERIN. 33

shore, 15 fathoms; bottom small gravel and shells, with
Cycloporus papillosa, Hyas coarctatus, Stenorhynchus
tenwirostris, Venus fasciata, Lissocardium norvegicum.

3. West of Fleshwick, further north, half a mile off
shore, 15 fathoms; bottom large shells, a little gravel,
with Pecten tigrinus, Venus casina, many common crabs.

4. One mile north of Fleshwick, half a mule off shore,
14 fathoms; bottom much fine gravel, with Pecten maxi-
mus, Trochus magus, Antedon rosacea.

5. Off the Cronk, a mile off shore, 14 fathoms; bottom
small gravel and some WMelobesia, with Tellina crassa
(alive), Thracia pretenuis. —

6. One mile further north, a mile off shore, 10 fathoms;
bottom Nullipores (Melobesia and Lithothamnion), with
compound ascidians.

7. West from South Barrule, a mile off shore, 12
fathoms; bottom Nullipores, with Antedon rosacea.

8. Off Niarbyl Point a mile out (several hauls), 12
fathoms; rough hard ground, with <Antedon rosacea,
Echinocardium flavescens.

9. Off Glen Meay, 4 miles out, 20 fathoms; bottom
“veamy”’ (sand and mud), with Ophiopholis aculeata,
Porama pulvillus.

10. Off Glen Meay, half a mile further north, 21
fathoms, with many Pecten opercularis, Cucumaria hynd-
mam, Ebalia tuberosa, Cellaria fistulosa, Scalpellwm
vulgare.

11. West of Contrary Head, 4 miles off, 18 fathoms ;
bottom Melobesia and stones, with Hugyra glutinans.

12. West of Contrary Head, 13 miles off, 13 fathoms ;
bottom muddy sand with some stones and many ophiu-
roids, with Cliona celata (massive form), Astarte sulcata,
Pecten maximus.

Il. March 25th. 1. Off Aldrick (south of Port Erin), a

34 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

mile out, 18 fathoms; bottom dead shells, shell sand, and
echinoderm spines, with Spatangus purpureus, Echino- .
cyamus pusillus, Porama pulvillus, Henricia sanguino-
lenta, Murex erinaceus, Xantho tuberculatus.

2. Off Kitterland, 14 miles out, 18 fathoms; bottom
dead shells, with Ascidia mentula, Cynthia morus. fae

3. North-west of Calf Sound, 2 to 3 miles. off, 19
fathoms; bottom sand and shells, with Palmipes placenta,
Lwidea ciliaris, Stichaster roseus, Thyone fusus and T.
raphanus, Cellaria fistulosa, Ascidia plebera, Polycarpa
comata.

4. North-west of Calf Island, 3 miles off, 20 fathoms ;
bottom sand and shell fragments, with Pectunculus gly-
cumeris, Modiola modiolus, Pecten maximus.

5. North-west of Burrow Rock, 3 to 4 miles off, 22
fathoms; bottom shells, with Pectwnculus glycimeris,
Lissocardium norvegicum, Pecten maximus.

6. North-west of Chicken Rock, 5 miles off, 25 fathoms;
bottom dead shells and some sand, with Sarcodictyon
catenata, Chetopterus sp., Hbala tuberosa, Ascidia
plebera.

7. One-and-a-half miles off Bradda Head, 18 fathoms;
bottom large shells and broken fragments, with Asterias
rubens (very large*), Porana pulvillus, Ciona intestinalis.

8. Iwo to three miles N.W. of Bradda Head, 21 |
fathoms ; bottom muddy sand, with a ophiuroids,
Cucumaria hyndmana.

9. Four miles N.W. of Bradda Head, 23-25 fathoms
(several hauls) ; bottom sandy mud, many ophiuroids. |

* The specimens we dredge are very much larger than those we find on
the rocks of the neighbouring shore. Are there two varieties in the species,
a smaller shore and a larger deep-water form, or do the indivi move
outwards from me shore as they grow older ?

MARINE BIOLOGICAL STATION AT PORT ERIN. a5

It. May 27th. The Committee hired the steam trawler
“Lady Loch,” and dredged the following localities :—

1. South-east of Calf Sound, a mile from Kitterland,
20 fathoms; bottom subangular gravel (? glacial material),
many ophiuroids and Buccinwm undatum, a few large
shells, Mytilus edulis, and Venus casina.

2. South-east of Calf Sound, half a mile further out, 19
fathoms; some coarse sand and broken shells, with the
subangular gravel (stones much incrusted), Spatangus
purpureus, many encrusting polyzoa, Venus, Trochus,
_Pecten, Serpula, Hchinus, and Lithothanuuon fragements.
38. South-east of Calf Sound, further on, 2 miles from
Kitterland, 20 fathoms; bottom white shelly (calcareous)
sand, mainly organic, lamellibranch and gastropod shells,
echinoderm spines and plates, Cellaria fistulosa and
Cellepora pumicosa, Pectunculus glycimeris, Zoophytes.

4. South-east of Spanish Head, 24 miles off, 20 fathoms;
bottom sand and broken shells; a few small stones—
Triassic sandstone, slate, and pebble of felsite. Annelids,
Phyllophorus drummondt.

5. South-east of Spanish Head, 3 miles off, 22 fathoms ;
bottom more shelly (fragments large), and a few small
pieces of slaty rocks.

6. Off the Chasms, half a mile out, 17 fathoms; bottom
muddy sand with much Lithothamnion and Melobesia, a
few shells and small stones, small sub-angular fragments
of slate, grit, Carboniferous limestone (with Productus),
and pebbles of coarse sandstone.

7. Off the Chasms, a mile out, 19 fathoms; bottom mud
and small gravel (subangular grit and granite), Hchino-
cardium cordatum and Hchinus remains, and some shells.

8. Off the Chasms, 2 miles out, 21 fathoms; mixed
bottom, sandy mud, small subangular stones and shell
fragments; with Pagurus prideauxu and Adamsia palliata,

36 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

9. South-east of the Old Mines, near Perwick Bay,
quarter mile to a mile off shore, 15 to 18 fathoms (two
hauls) ; bottom Nullipore and gravel (angular grit, slate,
vein-quartz) ; a few shell fragments with Hbala tuberosa,
Hyas coarctatus.

10. Off mouth of Perwick Bay, half a mile off, 12
fathoms ; bottom small gravel.

IV. July 8th. The Committee had the use of the
Lancashire Sea Fisheries steamer ‘‘ John Fell,’ and
dredged at the following localities :—

1. West of Dalby, 5 miles out, 30 fathoms; bottom
sandy mud, with Antennularia ramosa, Ophiura ciliaris
and O. albida, Pecten opercularis and P. pusio, Turritella
terebra, Scalpellum vulgare, Hyas coarctatus, Hupagurus
thompson, Hudorella nana, Gammaropsis (? n. sp.), both
the latter new to Britain, Ascidia virginea, and Hugyra
glutinans. ; |

2. Six-and-a-half miles west of Contrary Head (Peel)
38 fathoms; bottom fine mud, with Brissopsis lyrifera,
Lipobranchius jeffreysu, Hudorella nana.

3. Seven-and-a-half miles west of Niarbyl Point, 45
fathoms; bottom fine mud, with Calocaris macandree,
Gonoplax rhomboides, Harpyma levis (new to Britain).
Panthalis oerstedi, and fragments of a small Lumbri-
conereis.

4, Five-and-a-half miles west of Glen Meay, 34 fathoms;
bottom mud, many Twrritella terebra with Sagartia
herdmant.

5. Four-and-a-half miles west of the Cronk, 22 fathoms;
bottom broken shells and small stones, with many ophiu-
roids, Stichaster roseus, Hbalia tuberosa, Hurynome aspera,
Atelecyclus septemdentatus, many encrusting polyzoa
(twelve species identified), including Barentsia nodosa,
hydroids (fifteen species identified), including Dicoryne

MARINE BIOLOGICAL STATION AT PORT ERIN. ot

conferta, new to the district: also the cumacean Campy-

—laspis macrophthalma, new to Britain.

V. August 19th. The Committee hired the steam
trawler ‘‘ Lady Loch,” and dredged at the following
localities :—

1. North of Calf, close in, 14 fathoms, gravel, with
Clavelina lepadiformes, Asterias glacialis, many ophiu-
roids.

9. One-and-a-half miles north of Kitterland, 19 fathoms;
stones and mud, with Antennularia antennina and A.
ramosa, Sarcodictyon catenata, Chetopterus sp., Stichaster
roseus, Porama pulvillus, Hbalia tuberosa, Xantho tubercu-
latus, Ascidia mentula, A. plebeva, Corella parallelogramma,
Perophora listert, Forbesella tessellata, and Cynthia morus.

3. Two miles west of Bradda, 23 fathoms; Munida
bamffica, Ascidia mentula, Cynthia morus.

4. Mouth of Port Erin Bay (with small trawl), much
seaweed, with Ascidia albida (scabra), several common
species, such as Pagurus bernhardus and shrimps, some
young plaice and lemon sole.

VI. August 25th. In hired steam trawler ‘‘ Albatross.”’

1. Two miles north of the Calf, 22 fathoms; sand and
Pectunculus shells and Zoophytes, with Alcyonwum digi-
tatum, Sarcodictyon catenata (yellow variety), Campanu-
laria verticillata, Chetopterus sp.

9. Seven miles off outer end of Calf Id., 30 fathoms ;
dead Pectens and a few stones (up to 3 lbs.), in a little
mud, with Zoophytes, Cellaria fistulosa, Echinocardium
flavescens, Echinocyamus pusillus, Lyonsia norvegica.

8. Hight to nine miles west of Port Erin, 35 fathoms ;
bottom ‘“‘reamy,’ with Spatangus purpureus, Phascolo-
soma vulgare.

4. A mile further out, 48 fathoms; mud, (containing
some stones and dead shells), with Calocaris macandree,

38 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

with Triticella boeckii, Panthalis oerstedi, and Brissopsis
lyrifera. aa

5. Twelve miles off, 51 fathoms ; same mud and fauna.

6. Four miles west of Chicken Rock, 38 fathoms;
‘“reamvy’”’ and shells, Hwmenis jeffreysii, Thyone fusus,
Xantho tuberculatus, Lima elluptica, Pecten pusio.

7. Three miles west of Chicken Rock, 29 fathoms; large
dead shells, Crania anomala, Trochus millegranus, Corbula
gibba, Dentalium entale.

8. One-and-a-half miles west of Chicken Rock, 26
fathoms; shells, sand, and gravel (general colour yellowish
brown), several hauls, much material, Solaster endeca,
many ophiuroids, Hbalia tuberosa, Galathea, Aphrodite
aculeata, Dentalwum entale, Anonna ephippium, Pectun-
culus glycimeris, Venus casina, and V. gallina, Astarte
sulcata, Murex erinaceus, Polycarpa comata, Ascidia
mentula, and A. plebera, Corella parallelogranuna.

9. One mile off Calf, 25 fathoms; large stones, with fine
colonies of Sertularia abietina, Cynthia morus, Saxicava
rugosa burrowing in the limestone. |

10. Mouth of Port Erin Bay (small trawl) ; alge, with
common species of Stenorhynchus, Pagurus, Pandalus,
Crangon, Hydractuma, Natica, Lacuna.

VIL. September 30th. In hired steam trawler ‘‘ Lady
iioeh.

1. Five hauls were taken from 74 to 11 miles N.W.

Bradda Head, 47 to 60 fathoms; mud, with Twurritella . ,

shells; Brissopsis lyrifera, Panthalis oerstedi, Calocaris
macandee, and Triticella boeckw. Several of the Calo-
carts were females with eggs on the abdominal appendages.

2. From 8 in to 5 miles west of Dalby and Niarbyl,
about 20 fathoms; Sarcodictyon catenata, Porama pul-
villus, Palmipes placenta (a 6 rayed specimen), A glaophena
_tubulifera, Adamsia pallrata, Scalpellum vulgare, Hbala

MARINE BIOLOGICAL STATION AT PORT ERIN. 39

tuberosa, Lyonsia norvegica, Lima loscombu, Dentaluum
entale.

3. Off Fleshwick and round Bradda Head, half a mile
off shore (with small trawl), 18 fathoms; alge, with
common species of Hchinus, Asterias, Solaster, Stenor-
hynchus, Inachus, and Hyas.

It may be of some use to place on record the course of
procedure at each dredging station on these expeditions.
The plan for the day is arranged with the captain of the
steamer, and when the first locality is reached the spot is
determined on the chart, and the depth verified by casting
the lead. Then the dredge (measuring 2 feet 6 inches by
1 foot, and weighing from 380 to 40 lbs.) is sent down with
a tow-net tied on the line about two fathoms from the
dredge. Very often a smaller dredge with a bag of
cheese-cloth is sent over on the other side of the ship.
One or more surface tow-nets are also put out. The
tow-nets, both surface and deep, are looked after by Mr.
I. C. Thompson, who, after hauling them, first turns out
their contents into a clear glass jar of sea-water, and then,
after noting the general character of the catch and any
specially conspicuous forms, strains off the water through
a small bag made of very fine miller’s silk, and transfers
the ‘‘plankton”’ left adhering to the silk into a tube
containing a special preservative fluid formed of spirit,
glycerine, and water.

When the dredge is brought up it is emptied on deck,
and after a note of the general character of the deposit
and assemblage of animals has been taken, any specially
large or rare specimens are picked out and transferred to
buckets or jars of sea-water, or to store-bottles of spirit.
Then the heap is spread out so as to form a layer not
more than one or two inches in depth, and one or two

40 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

members of the Committee (Professor Herdman and
another) now settle down beside it to pass the entire
mass in review inch by inch, working it across a small
space of bare deck and turning over every shell, stone and
specimen with an iron spoon, so as to ensure that nothing
escapes observation and due record in the note-book. In
the meantime the contents of the bottom tow-net have
been dealt with by Mr. Thompson, and the apparatus has
been lowered for a second haul, or the vessel is steaming
on to a new locality. Then Professor Herdman selects a
fair sample of the deposit for preservation (for the Geo-
logical Survey) in a small canvas bag (10 by 5 inches),
care being taken to include some of the characteristic
bottom animals—shells, ophiuroids, polyzoa, &c. After
this sample has been removed, and any special animals
required have been picked out and put into store-bottles,
the whole of the remainder of the haul is passed gradually
through our set of three sieves (meshes 2 inch, + inch,
and 4+ inch respectively), which work up and down in a
tall iron cylinder filled with sea-water. The sieves are
disconnected and examined at intervals, and in this way
many of the smaller animals of all groups are detected
and picked out. Finally, the water in which the sieves
have been plunging is all strained by Mr. Thompson
through his fine silk net, and in this way many of the
rarer bottom Copepoda are obtained, while the finer sandy
and muddy deposits retained by the finest sieve or in the
bottom of the cylinder are packed in canvas bags by Mr.
Alfred Leicester for further examination at home. These
contain, of course, many minute Mollusca, Ostracoda, and
Foraminifera. By the time all these processes have been
completed the dredge has usually been hauled again, and
a fresh heap is lying on the deck awaiting investigation.
On a successful trip the members of the party, on an

MARINE BIOLOGICAL STATION AT PORT ERIN. 41

average four to six in number, are kept constantly occu-
pied, each man at his own work, from the commencement
of the first haul till the steamer is turned homewards, and
after that the packing and labelling of specimens fill up
the time until land is reached.

ADDITIONS TO THE FAUNA.

As most of the expeditions took place round the Isle of
Man, the material was generally brought back to the Port
Erin Biological Station, and sorted out into groups in the
laboratory there, and then sent to the specialists. Taking
the groups in zoological order, the most notable additions
as the result of this year’s work have been—

SPponcES.—Dr. R. Hanitsch reports that the only actual
additions to our sponge fauna made during the last few
months are (1) Letosella (Spongionella) pulchella, Sowerby,
which was dredged on May 14th, 1894, at 14 miles N. by
W. from the Liverpool N.W. Lightship. This species
was previously known from the coast of Durham, the
Skerries, Shetland, the west coast of Ireland, the east
coast of Greenland, and the North Pacific. (2) Myzilla
wrregularis, B., found at Port Erin and Fleshwick, at low
tide, in August and September, 1894. This species has
previously been recorded only from the Diamond ground
off Hastings. We have also found at Port St. Mary a
very fine specimen of Lewcandra gosset, consisting of at
least 16 ‘‘ persons,’ while Haeckel describes the species
as having 2 to 5, rarely 4 to 8 persons.

We are indebted for a list of the HypRorp ZOOoPHYTES
and Pouyzoa which we have collected to Miss L. R.
Thornely, who has proved that the Lafoéa pigmea of Alder
possesses an operculum, and therefore belongs to the
genus Calycella, and also has gonothecwe, which were
previously unknown. ‘The total number of species of

42 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

hydroids in our area is now ninety, and one of the dredg-
ing expeditions has given us interesting additions to
our fauna in Dicoryne conferta, which was growing on an.
Aporrhais shell, and was only known previously from
Cullercoats, Orkney, and Shetland; and Syncoryne exumia,
with numerous sacks or “‘ galls”’ containing young Pycno-
gonids. Of polyzoa 123 species and 15 varieties have
now been recorded. The most recent finds are Crisia
ramosa, which was recently described by Harmer from
Plymouth, and which we find also at Port Erin, and
Microporella invpressa, var. a., found by Miss Thornely at
Port Erin in July.

Mr. E. T. Browne has, during some visits to Port Erin,
paid special attention to the Mmpus#, and has drawn up
a list of about twenty species, one of which, at least,
Anwlicodon fritillaria, has not previously been recorded
for British seas. The most interesting capture which he has
made consists of several specimens of a beautiful Siphono- —
phore, which appears to be a species of Halistemma, cer- —
tainly new to the district and probably to the British fauna.
Professor M‘Intosh recorded a Halistemma in 1891 from
St. Andrew’s Bay, but our Port Erin species is not the
same. Mr. Browne describes the colony as being about
25 mm. in length when expanded; it has a pneumato-
phore, about 6 large nectocalyces, about 4 polyps, with
hydrophyllia, and long tentacles armed with clusters of
nematocysts. ‘The polypes and pneumatophore are of a
deep reddish colour. ‘The colonies were found in April,
1894.

The yellow variety (?) of Sarcodictyon (Clavularia)
catenata, of which we dredged several colonies on August
25th, off the north-west of the Calf Is., in 22 fathoms, is
an interesting addition to our fauna. It has only been
found before in Loch Fyne and at two other spots on the

vy

MARINE BIOLOGICAL STATION AT PORT ERIN. 43

west coast of Scotland. Some sections which I have
made recently of these specimens show that the colonies
were then (August) probably ready for reproduction, as
the polypes contain mature ova. This is probably the
Sarcodictyon agglomeratwm of Forbes and Goodsir ; a note
on the subject will shortly be laid before the Biological
Society.

Amongst Worms new to the record are the Turbellaria
Fecampia (the pear-shaped white cocoons of this form are
not uncommon on stones in pools at Port Erin), and
Stylocoplana maculata (identified by Mr. Gamble), and
the annelid Amblyosyllis (Gattiola) spectabilis (Johnston)
collected at Port Erin by Mr. Beaumont.

Professor G. 8. Brady has kindly examined a number
of gatherings of OsTRAcoDA, from dredged material, taken
from 10 to over 40 fathoms, and reports the following
species :—

Pontocypris trigonella, G. O. Sars; P. mytilovdes, Nor-
mame P. serrulata, G. O.S.; Bardia inflata, Norman;

WOythere jones, Baird; C. tuberculata, G. O. Sars; C.

pao, ©. B.; C. fmmarcinca, G. O. 8:; C. confusa,
B. and N.; C. concinna, Jones; C. dwnelmensis, Norman ;
C. antiquata, Baird; C. emaciata, G. 8. Brady; C. con-
pera, baird; C. villosa, G. O. 8.; Hucythere argus,
G.O.S. Krithe bartonensis, Jones; Loxoconcha wmpressa,
Baird; L. guttata, Norman; L. tamarindus, Jones; L.
pusilla, G. 8. B.; L. multifora, Norman; Cytherura
cornuta, G. 8. Brady; C. angulata, G.5.B.; C. cellulosa,
merman; C. striata, G. O. Sars; C. sella, G..O. Sars;
Pseudocythere caudata, G. O. Sars; Cytheropteron latis-
suemum, Norman; C. pyramidale, G. 8. B.; C. alatum,
G. O. Sars; Sclerochilus contortus, Norman; Paradoxo-
aroma normam, G. S. B.; P. ensiforme, G. 8. B.; P.
variabile, Baird; P. hibernica, G. S. B.; Philomedes inter-

44 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

puncta, Baird; Cytheridea papillosa, Bosquet ; C. punctil-
lata, G. 8. B.; Bythocythere acuta, Norman; B. constricta,
G.O.8.; B. turgida, G. O. Sars; Macherina tenwissima,
Norman.

The common Lozoconcha impressa is extremely abundant
sometimes on weed brought into the laboratory at Port
Erin from the shore pools. About the Cytheropteron
pyramidale noted above, Prof. Brady writes as follows :—
“T find that it must be referred to C. pyramidale, a
species of which there has been hitherto no undoubted
British record. It comes very near to C. latissemum
which, on our east coast, is a common species, but is
not found on the west. In a gathering from 14 miles
N. by W. of the Liverpool Lightship, C. pyramidale is
quite the prevailing form. This 1s very interesting. I do

not think I have seen it in any of the Isle of Man
- gatherings.” |

In regard to the CopEpopa Mr. I. C. Thompson has
drawn up a general report upon the additions to our
knowledge of the group (see p. 46); while Mr. Andrew
Scott, ‘‘ Fisheries’’ Assistant at University College, has
suppled the following notes upon some new species of
Ectinosoma and other Copepoda, at which he has been ©
specially working :—

‘“« Longipedia nunor 'T. & A. Scott.—A few specimens
of this species were collected by hand-net in the rock-
pools at. Hilbre Islandin March. It is easily distinguished
from LZ coronata (Claus) by its much smaller size. |

‘““ Hctinosoma normam, n. sp. (LT. & A. 8.).—Several
specimens of this Hctinosoma were obtained in material
from Barrow Channel, collected by Professor Herdman in
May, and it was also found by Mr. Thompson at Port
Erin. When fresh this species has a brilliant red spot on
the lower angles of the cephalothorax, and in this respect
it agrees with EH. erythrops, Brady.

:
“A
J
=
;

MARINE BIOLOGICAL STATION AT PORT ERIN. 45

““ Hictinosoma gracile, n. sp. (T. & A. §.).—One or two
specimens of this species were obtained among dredged
material collected at Port Erin by Professor Herdman,
Easter, 1894.

‘“‘ Hctinosoma pygmeum, n. sp. (T. & A. 8.).—This
species was obtained from the same material as the last,
and is the smallest Hctinosoma known to us; it measures
only = of an inch (33 mm.)

Secumosoma herdmant, n. sp. (T. & A- S.).—One
specimen of this was found by Mr. Thompson at Port
Erin.

“ Bradya mimor, n. sp. (T. & A. 8.).*—A few specimens
of this new Bradya were obtained in rock-pools at Hilbre
Island, along with Longipedia minor.

“ Dactylopus rostratus, T. Scott.—A single specimen
was obtained among some dredged material collected at
Port Erin by Professor Herdman, at Easter, 1894.

“ Pseudanthessius sawvaget, Canu.—A few specimens
were obtained by washing a number of Spatangus pur-
pureus, which were trawled in the central area, 21 miles
W.N.W. from Morecambe Bay Lightship, on April 38rd.
This rare species was only added to the British fauna last
year, when it was found in the Firth of Forth, and the
present is the second time it has been observed in the
British area.”’

Lichomolgus (Doridicola) agilis (Leydig), was found in
the bottom tow-net, Morecambe Bay, May, 1894. Dvo-
saccus propinquus (T. & A. 8.), Amewra exiqgua, 'T. Scott,
A. longiremis, 'T. Scott, Laophonte inopinata, T. Scott,
Pseudowestwoodia pygmea, TV. & A. §., and possibly a
new Laophonte, and one or two other doubtful species

* The above Eetinosomas and Bradya are figured and described in a revision

of the British species of Copepoda belonging to the two genera Letinosoma
and Bradya, by T. and A. Scott, which is to be published at an early date

46 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

were all obtained from washings from sponges collected
by Dr. Hanitsch at Port Erin in August, 1894.

Mr. Thompson reports as follows :—

‘““In addition to those mentioned by Mr. Scott, seven
species of Copepoda, new to the district, have been re-
corded during the past year, viz., Cyclops magnoctavus,
Cyclops ewarti, Centropages typicus, Ameira longicaudata,
Acontiophorus elongatus, Cancerilla tubulata, and also
one species new to science, viz., Pseudocyclopia stephoides,
n. sp. This crustacean has just been described, and its
description and figure will be shortly published in the
‘Transactions of the Liverpool Biological Society.’ It
combines some of the characters of the genus Stephos
with those of Psewdocyclopia, the latter predominating
sufficiently to determine its position in that genus.

‘“ Surface tow-nets have been continually employed
during the several marine expeditions undertaken by the |
Committee, also tow-nets attached to the rope a few
fathoms above the dredge. The latter device has proved
a success, collecting some good species of Copepoda, as
well as Cumacea and Amphipoda, which are seldom or
never obtained on the surface. Amongst the Copepoda
thus obtained were several specimens of Psewdocalanus
armatus, found along with a shoal of Psewdocalanus elon-
gatus. A widely extending shoal of Anomalocera pater-
sonit was observed off the Isle of Man in May, the only
occasion on which we have taken this species during the
year. On several occasions during the past summer we
have taken Centropages typicus in fair quantity, a species
which has never occurred before in our district, although
carefully looked for, and not a rare species elsewhere.
On several occasions, notably in the early part of June,
the surface organisms have been singularly scarce.

‘‘ Special care has been taken to wash and sieve through

MARINE BIOLOGICAL STATION AT PORT ERIN. AT

fine silk as much as possible of the material brought up
by the dredge during marine expeditions, and it is by this
means that several of the above-mentioned Copepoda new
to the district have been obtained, as well as the new
species Pseuwdocyclopia stephoides. large quantities of
ophiuroids, chiefly Ophiocoma mgra and Ophlothria
fragilis, were amongst the dredged material, and it 1s pro-
bably from one or other of these that the two specimens
of Cancerilla tubulata, Dalyell, a male and female, were
taken, as the species is parasitic on ophiuroids. The first
record of this rare copepod occurs in Dalyell’s ‘“‘ Powers
of the Creator,’ 1851, and it has since been taken by
Mr. Gamble at Plymouth, and off the French coast, but
not before in our district. Cyclops magnoctavus, Cragin,
was found along with quantities of T’emorella affims and
Tachidius brevicornis in tow-nettings taken by Mr. Ascroft
in low-water marine pools at Lytham: these being
brackish species, it is probable that a considerable
amount of fresh-water finds its way into the Lytham
pools. Cyclops ewarti, Brady, although first taken in the
Forth estuary, was suspected by Brady to have a fresh-
_ water origin. Ours are evidently strictly marine, two
specimens, both males, having been dredged at 20 fathoms
by Mr. Thompson at Port Erin.”

Mr. A. O. Walker reports as follows upon the HIGHER
CRUSTACEA :—“‘ Collections have been examined from the
following places, viz.—

1. Off Port Evin at various points, dredged in (usually)
10 to 50 fathoms by Professor Herdman and Mr. I. C.
Thompson, in March, July, August, and September, 1894.

2. Off the Little Orme, North Wales, 5 to 10 fathoms
(dredged by A. O. Walker).

3. In the Menai Straits, near the Suspension Bridge
(both above and below), on April 2nd and May 3lst, 1894
(dredged by A. O. Walker).

48 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

The additions made to the list published in last year’s
report are marked with an asterisk. |

PODOPHTHALMATA.—Gonoplax rhomboides, Linn., one
specimen on July 7th, 1894, on mud, 45 fathoms; 7 miles
W. of Niarbyl, Isle of Man.

* Pisa biaculeata, Mont., off Port Erin, Easter, 1894.

Palemonetes varvans, Leach, in a small pool by the
Afonganol, Colwyn Bay, in company with Neomysis vul-
garis. The pool had probably been filled by a combina-
tion of flood in the httle river and a high tide, but seemed
to have been long cut off. The pool was full of Ruppia
marituma. The Palemonetes were 40 mm. long, and
females had ova in the pouches; the Neomysis, on the
other hand, were small, females with ova being only 14
mm. long.

CumMacHA. — * Nannastacus unguiculatus, Bate, one
specimen from Menai Straits.

*Campylaspis macrophthalma, G. O. Sars, one female —
from 44 miles west of the Cronk, July 8th, 1894, 22
fathoms. Thisis a Mediterranean species, new to Britain.

*Petalosarsia declivis, Sars, 8 miles W. of Fleshwick
Bay, 33 fathoms; from bottom tow-net at 14 miles N.W. |
of Liverpool N.W. Lightship (A. Scott). This species,
first recognised in our district by Mr. Scott, is only known
elsewhere in British seas from the Firth of Forth and the
Moray Firth. It is therefore an addition to the west
coast fauna.

*Cuma puichella, Sars, re off Little Orme, in Sept.

Tsopopa and AMPHIPODA.—Cymodoce truncata, Leach,
off Port Erin. .

*Hudorella nana (Sars), in mud, at 30 and at 38
fathoms, West of Dalby and Peel, on July 8th, 1894 (new
to Britain). This agrees with Sars’ description, but a
re-examination of specimens reported as EH. truncatula

MARINE BIOLOGICAL STATION AT PORT ERIN. 49

from Puffin Id., shows that the teeth on the lower anterior
margin of the cephalothorax differ from both species, so
that this may be a variable character. If so, H. nana can
hardly be separated from H. truncatula, Bate.

*Nannonyx spinimanus, n. sp., Menai Straits, differs
from the only other known species in the following points:
Anterior coxal plates about the same depth as the body;
eyes very large, dark brown; propodos of first gnathopod
with five or six strong denticles on the hind margin; third
joint of last pereiopods but little expanded behind; maxil-
lipedes also different.

*Socarnes erythrophthalmus, Robertson; Port Erin
15 to 20 fathoms, March 24th, 1894; Menai Straits.

Urothoé brevicornis, Bate; off Kitterland, 20 fms., male.

Phoxocephalus fultom, Scott; Port Erin, 15 to 20
fathoms; Menai Straits.

*Harpuua levis, Sars, two specimens in mud, from 45
fathoms; 74 miles west of Niarbyl, on July 8th (new to
Britain).

*Gammaropsis (? n. sp.), In sandy mud, from 380
fathoms, W. of Dalby, on July 8th. This is probably a
new species near G. melanops (Sars), but differing in the
form of the propodos of the first gnathopods and in the
secondary appendage of the upper antenne, which is
three jointed, the last joint being minute.

Leucothoé lilljeborgu, Boeck; Port Erin, 15 to 20 fms.

*Iphivmedia nunuta, Sars; Colwyn Bay, Port Erin.

Mera batet, Norman; Port Erin.

Leptocheirus pectinatus, Norman; Port Erin, Menai Str.

Autonoe longipes, Lilljeborg; Menai Bridge.

Janassa capillata, Rathke; Port Erin.

Colomastia pusilla, Grube; Menai Straits.

Corophium bonellu, M. Kdw.; Little Orme, Port Erin.

There still remains a quantity of material to be ex-
amined.”’

50 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. —

My. Alfred Leicester, of Southport, who has taken part
in most of the expeditions, and has collected and identified
the Mollusca, reports that the year’s work has added fifty-
one fresh records to the lists for the southern part of the
Isle of Man, and that of these the following nine are new
to our district of the Irish Sea:—Cardium minumun,
Phil., Psammobia vespertina, Chem., Scrobicularia mtida,
Mull., Chiton marginatus, Penn., Propilidiwm ancyloides,
~ Forb., Riassoa inconspicua, Ald., Cecwm trachea, Mont.,
Aclis gulsone, Ch., and Philine angulata, Jef.

Finally two additions have been made to our list of
local fishes, viz., Zeugopterus unvmaculatus, four specimens
trawled 10O—12 miles west from Morecambe Bay Lightship
in May, depth 23 fathoms; and Gobius pictus, Malm,
caught by Mr. Walker in shore pools at Colwyn Bay.

THE SUBMARINE DEPOSITS.

Turning now to the submarine deposits, the determina-
tion and distribution of which the Committee feel to be a
very important part of their work, it is still too soon to
attempt anything like a detailed account of the floor of the
Irish Sea, but still sufficient observations have perhaps
been made to warrant the following preliminary account.
The accompanying chart (Plate I.) shows the zones of
depths in the district, O—10 fathoms, 10—20 fathoms,
20—50 fathoms, and upwards oi 50 fathoms, being
separated from one another. At those places where the
Committee have obtained samples of the bottom, conven-
tional symbols are placed on the map* indicating, © stones,
A shells, = mud, [x| sand, x nullipore deposits (Melobesia

and Lithothamnon), and On shell concretions. The
chief conclusions we have arrived at so far are :—

* One mark frequently stands for a number of different dredgings in the
same neighbourhood.

MARINE BIOLOGICAL STATION AT PORT ERIN. 51

1. The most extensive shallow-water deposit is sand.
In most localities along the coast of Lancashire, Cheshire,
and North Wales, from the sea-shore out to the 10-fathom
contour, the bottom is formed of more or less pure quartz
sand. Occasionally in spots there are local patches of
stones, of shells, or of mud; but these can generally be
accounted for by tidal or estuarine currents, by the entrance
of fresh-water streams carrying down alluvium, or by the
presence of littoral or sub-littoral boulder clay. These
spots are all, however, of small area, and the great extent
of the bottom down to 10 fathoms is sand.

2. Further out, however, between 10 and 20 fathoms,
the sand becomes greatly mixed with mud, and much
diversified by large tracts of shelly deposits or by patches
of gravel, and the fauna on the bottom also becomes much
more abundant. In some spots, at about 20 fathoms, it
is made up over considerable areas almost entirely of
ophiuroids (Ophiocoma nigra and Ophiothria fragilis),
which fill the dredge haul after haul. At two localities
off the Isle of Man, viz., along the east coast from Clay
Head to St. Ann’s Head, and off the west coast between
Contrary Head and Niarbyl, at depths between 10 and 20
fathoms, are great nullipore deposits formed of Melobesia
and Lithothamnion, which have a most characteristic
appearance, smell, and fauna.

This area of the sea-bottom, from 10 to 20 fathoms,
extends across from the north of Lancashire to the Isle of
Man, so that opposite Barrow, for example, there is a
wide extent of about 50 miles in length of sea-floor at
depths of not more than 15 or 16 fathoms. ‘The Isle of
Man is connected with England by this plateau, and is
separated from Ireland by deep water.

3. Depths of over 20 fathoms are only found to the
west, north, and south of the Isle of Man; and depths of

52 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

from 20 to 50 fathoms give us the most varied bottom
deposits and the richest fauna. As a rule, the sand is
more or less mixed with mud, and as the bottom goes —
deeper the amount of mud gets greater. When there is a
considerable admixture of mud with coarse sand, that
forms what is known to the trawlers asa ‘‘reamy’’ bottom:
and that is the ground upon which the sole and some
other fish are generally found spawning.

Shells and other hard parts of animals play an important
part in the deposits at depths of about 20 fathoms and
upwards. In places the dredge comes up filled with
Pecten shells, dead and alive, chiefly P. opercularis and
~P. maximus. At other places the deposit is practically
composed of the shells of Pectunculus glycumeris. These
and other shell beds form a rich collecting ground to the
naturalist, as they support an abundant and varied fauna.
Loophytes and polyzoa are attached to the shells, and
these serve as shelter for nudibranchs and other small
mollusca, worms, and ascidians. On the whole the hete-
rogeneous deposits support a richer fauna than do the
homogeneous deposits, such as sand or mud, and it is
chiefly in the zone of depth we are now considering that
the heterogeneous deposits occur.

4. The depths over 50 fathoms contain a pure dark
bluish grey mud, which is very tenacious, and sets when
dried into a firm clay. This is abominable stuff to dredge
in and to work with on deck. It clings to everything that
touches it; it is almost impossible to see what is in it, and
to get the animals out of it uninjured; it is too solid for
the sieves, and the hose can be played upon masses of it

almost indefinitely without dissolving it. The fauna of — !

this zone is, in our district, quite peculiar and character-
istic. In its shallower parts, about 50 fathoms, it contains
great numbers of living and dead Turritella terebra, upon

MARINE BIOLOGICAL STATION AT PORT ERIN. Do

many of which are attached one, two, or three specimens
of the little red anemone, Sagartia herdmam, Haddon.
In its deeper parts, up to 80 fathoms, are found Calocaris
macandree, Hyalinecia tubicola, a small Lumbriconereis,
Panthalis oerstedi, Lipobranchius jeffreysu, Brissopsis
lyrifera, Amphiura chiaju, and Isocardia cor. Numbers
of large sausage-like muddy tubes, formed of stratified
layers of interlacing threads of mucus in which the mud
particles are closely entangled, are brought up in the
dredge. These we have now proved to be the tubes of
Panthalis oerstedi, and the living annelid has several
times been found in the tubes, but most of those we
dredge up are empty, and the tubes are certainly far more
numerous than the worms. Possibly the explanation is
that the Panthalis forms a tube as it lies in the mud, and
then when it moves away leaves its tube behind it (one
can scarcely imagine the animal dragging such a tube
through this tenaceous deposit), and after a time forms
another in a new situation.*

These are the leading conclusions we have come to so
far in regard to the distribution of sub marine deposits in
our area. ‘T'wo further questions now present themselves ;
first, the biological one—the effect upon the fauna; and
secondly, the geological one—the origin of the deposits.
In regard to the importance of the nature of the bottom
to the animals living upon it there can be no doubt.
Probably the nature of the deposit is the most important
of the various factors that determine the distribution of

* This suggested explanation was written in June, and was read in August
before Section D of the Brit. Assoc. at Oxford. The work of Mr. Watson, at
Port Erin at the end of August has pretty well established its correctness, as
he watched a Pantha/is, in a tank, desert its old tube and form a new one
from mucus threads and mud particles (see p. 61). It must be remembered
also that some of the empty tubes doubtless belong to dead worms,

54 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

animals over the sea-bottom within one zoological area.
It is certainly more important than mere depth; a muddy
bottom will support a similar fauna at 10 fathoms in one
place and at 50 fathoms in another. Probably the most
important influence in the environment of a lower animal
is 1ts food, and once beyond the narrow sub-littoral zone
in which alge flourish—and to which, of course, certain
phytivorous animals must be restricted—it is probably
chiefly the nature of the bottom which determines the
food.* Many animals feed upon the deposit, others
browse upon the polyzoa and zoophytes which can only
attach themselves and grow where there are sufficiently
large objects, such as shell valves, from which they can
get the necessary stability; while others, again, feed upon
their neighbours, which subsist on the deposit or are -
attracted by the zoophytes, &c.; for example, soles are
frequently caught upon ground (known to fishermen as
“sole ground’’) where Flustra folvacea lives in abundance,
and the probable connection is that the fish are dependent
upon the numerous amphipoda and other small animals
which frequent the tufts of Flustra. The same locality
may vary so much from time to time in the temperature,
the salinity, and the transparency of the water, that it is
probable that none of these factors—so long as the varia-
tions do not exceed certain limits—have so much influence
upon the fauna as the nature of the deposit has. It is
therefore quite to be expected that the fauna should vary
from place to place with the nature of the bottom, and
that is what we have observed frequently in our work
round the Isle of Man. In practically the same water,

* The only food supply quite independent of the bottom is dead plankton,
from the water above, which may reach the bottom uneaten ; and possibly a
small amount of decayed vegetation and other organic matter brought down
by rivers from the land, and some of which may reach the sea-bottom,

MARINE BIOLOGICAL STATION AT PORT ERIN. 55

identical in temperature, salinity, and transparency, at the
same depth, with, so far as one can see, all the other
surrounding conditions the same, the fauna varies from
place to place with changes in the bottom—maud, sand,
nullipores, and shell beds, all have their characteristic
assemblages of animals.

As to the further, and very important, question of the
origin of the deposits, that is to a great extent a purely
geological inquiry, and one which cannot, until we have
accumulated a much larger series of observations, be fully
discussed; but there are a few matters which may be briefly
pointed out as giving some idea of the range and bearing
of the question.

1. It is necessary to make a most careful examination
of the deposits. For example, all muds are not the same
in origin. A deposit of mud may he due to the presence
of an eddy or asheltered corner in which the finer particles
suspended in the water are able to sink, or it may be due
to the wearing away of a limestone beach, or to quantities
of alluvium brought down by a stream from the land, or
to the presence of a submerged bed of boulder clay, or,
finally, in some places, to the sewage and refuse from
coast towns.

2. Ihave kept in view the possibility of some correlation
between the geological formations along the beach and
the sub-marine deposits lying off the shore. There is no
doubt that the nature of the rock forming the shore has a
great influence upon the marine fauna, and has sometimes
some effect upon the neighbouring deposits. For example,
the contrast between the deposits lying off the two pro-
minent headlands, the Great Orme, in North Wales, and
Bradda Head, in the Isle of Man, is well marked. ‘The
Great Orme is composed of mountain limestone, and the
result of its weathering and erosion is that large blocks

56 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

are found lying scattered outside its base on the fine sand; —
but there is no deposit of smaller stones, gravel, and
resulting sand farther out, probably because in the wear-
ing of the rock and large detached blocks by the sea a
great deal is removed in solution and the rest in suspension
as very fine mud—this we have found to be the case round

Puffin Island, which is also mountain limestone. Bradda q

Head, on the other hand, is a schistose metamorphic
silurian rock, which breaks up into large fragments, and
these into smaller, and so forms deposits of dark slatey
more or less angular gravel, and then very coarse sand,
extending for some way out from the foot of the cliff.

The influence of the shore rocks upon the littoral fauna
is an important subject upon which we have accumulated
some observations; but the matter requires further work _
and detailed discussion, and must be left over for a future
report. |

3. Probably the great bulk of the silicious sand which
forms so large a part of the floor of our sea is derived proxi-
mately—whatever may have been its ultimate source*—
from the great deposits of drift which were formed in the
neighbourhood during the Glacial period, and large tracts
of which may since have been broken up by the sea.

4. As examples of a few peculiar and specially note-
worthy deposits which are not simply “ terrigenous”’ in
their origin, the following may be mentioned :—

South-east of the Calf Sound, about two miles out, at a
depth of 20 fathoms, there is a white shelly sand which -
seems to be almost wholly composed of animal remains.
There are broken fragments of the lamellibranchs Pecten,
Anomia, Pectunculus, Mactra, Venus, and Mytilus, of the
gastropods Cyprea, Buccinum, Hmarginula, Purpura, and

_* Probably to a great extent, Triassic sandstones. ~

MARINE BIOLOGICAL STATION AT PORT ERIN. 57

Trochus, of various calcareous polyzoa, such as Cellaria

fistulosa, Cellepora punvicosa, and lepralids, of Balanus
and Serpula, and of various echinoderm plates and spines,
and the whole shells of Hchinocyamus pusillus. The
deposit, when it comes up in the dredge, is of a gleaming
whiteness, and has a very characteristic appearance. Such
a deposit as this would form a rock almost wholly made
up of fossils, and might compare well with some Tertiary
fossiliferous deposits, such as the Coralline Crag.

A little further north, along the east coast of the Isle of
Man, at about a corresponding depth and distance from
land, we meet with a purely vegetal deposit formed of the
nullipores Lithothamnion and Melobesia. On the other
side of the island, again, between Port Erin and the Calf,
at a depth of 18 fathoms, there is a tract of sea-bottom
which, when brought up on deck, looks, at the first glance,
like a peculiarly fibrous sand, but a closer examination
shows that it is entirely composed of the comminuted
plates, and especially the spines of echinids, chiefly
Spatangus. I do not remember to have met with a
reference to material such as this either amongst recent
or fossil deposits.

The remarkable deposit of cemented shells, which was
described in last year’s report, from a locality half-way
between the Calf and Holyhead, is also found off King
William’s Bank, between Ramsay and St. Bees. Plate
II. represents a piece of this material.

The variety that is noticed in sub-marine deposits round
the Isle of Man, from depths of 15 to 35 fathoms, as
brought up in the dredge, is very striking. It is remark-
able how differing proportions in the mixtures of sand,
gravel, and shells give rise to very different colours and
general appearance in the mass. As seen, when tumbled
out of the dredge on to the deck, some deposits are white,

58 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

some yellow, some grey, some reddish, of various tints
from pink to ruddy brown, and others darker, of all
shades of brown and dark grey. It is curious how, even
in a composite deposit made up of many different con-
stituents, there is usually a prevailing tint; for example,
the bottom at one spot, although composed of mud, sand,
nullipores, shells, and stones, is distinctly of a rich ruddy
brown tint. The importance of this presence of prevailing
colours in the various sub-marine deposits is obvious in
its bearing upon the colours and habits of animals.

. Sample bags of all the more important sub-marine
deposits we have come upon have been sent, at Sir
Archibald Geikie’s request, to the Museum of the Geo-
logical Survey in Jermyn Street. They are being examined
there by Mr. Clement Reid, F.G.S., who writes the follow-
ing preliminary note in regard to them :—

‘“On comparing these samples with British deposits of
Tertiary date, one finds a marked difference in lithological
character. Dredgings from the Irish Sea, and also from
the North Sea, are characterised by a much coarser and
more gravelly texture than one would expect at such
depths—coarser, in fact, than one finds in Pliocene
deposits yielding a similar fauna, indicating similar or
even smaller depths. A glance at these dredgings shows
the reason for this, for they are largely composed of
unworn or little worn fragments of rock, often entirely
incrusted by organic growth. The stones evidently have
not been transported far by water, or they would be
well rounded, like the pebbles found in our Hocene beds.
The incrusting organisms show also that the fragments
have lain undisturbed on the sea-bed, yet they have often
been derived from far-distant sources. Though no Glacial
striz were observed, and no undoubted sub-fossil Arctic
shells have yet been found at these localities, yet there

MARINE BIOLOGICAL STATION AT PORT ERIN. bg

seems little doubt that the bulk of the material on the
sea-bottom over this area has been derived from the
breaking up of pre-existing Glacial deposits. This may
occur at a depth of several fathoms through the gradual
washing away of the muddy and sandy matrix of a boulder
clay or Glacial gravel. Coarse gravel is thus caused to
accumulate at a spot where the currents may be too
feeble to transport anything but sand.

“This sub-marine origin of angular gravel deposits
should not be forgotten, for it affects the lithological
character of the sea-bottom over most of the area which
was formerly glaciated, even as far south as Cornwall.
On the other hand, it does not affect, except to a small
extent, the sea-bed beyond the former limit of the ice, and
it does not affect pre-Glacial deposits. Thus we must
always expect to find at similar depths the same fauna
associated with deposits of finer texture as soon as we
leave the glaciated area, or when we go back into Ter-
tiary times.

“Tt is also worth noting that the occurrence of a stony
bottom at 20 or 30 fathoms—where normally there would
be no deposit coarser than sand—will probably lead to a
disproportionate increase of all incrusting organisms, and
of all organisms needing a solid base. This has certainly
taken place, as anyone studying our shoal-water Tertiary
deposits will have observed. They contain few stones,
and though each stone or dead shell may be covered with
incrusting organisms, yet the relative proportion of these
to the free forms is far smaller than seems commonly to
be the case in the seas that now wash our shores. ‘The
sole exception to this rule among the British Tertiary
strata is found in the Coralline Crag, in which the con-
temporaneous consolidation of the limestone was sufficient
to provide the necessary solid base for the incrusting and
fixed organisms so abundant in that deposit.”

60 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

In conclusion, it is clear that this investigation of our
modern sub-marine deposits, their distribution, nature,
origin, and associated fauna, has geological applications,
and that our results may be of some importance, e.g., to
paleontologists, in aiding them to determine the conditions
under which the fauna of a particular horizon probably
existed in the past; but, from our point of view, the matter
is a purely Biological one. We consider it of primary
importance, in studying the distribution of the marine
animals in our district, to investigate as minutely as
possible their environment, and that not merely because
it gives us some of the factors and possibly the explana-
_ tion of the distribution, but also on account of the light it
may throw upon the habits, variations, and other important
characteristics of the species.

OTHER FAUNISTIC WORK.

In addition to the larger dredging expeditions (see p.32),
a good deal of work has been carried on during the year
from small boats in Port Erin Bay, and by shore-collect-
ing at Port Erin, Port St. Mary, Perwick, Fleshwick, and
the Calf Sound.

One interesting piece of work carried out in the Aquar-
ium, was Mr. Arnold T. Watson’s observations on the
tube building of Panthalis oerstedi, a Polynoid worm.
As is stated above (p.53), in the deep water off Port Erin
we dredge in the mud large thick-walled tubes, which are
sometimes associated with Panthalis, and which we sup-
posed to be formed by that annelid. However, some
doubt had been expressed on the matter, and a leading
authority on the Annelida gave it as his opinion that the
connection between Panthalis and the tubes had never
been clearly established. Consequently, I suggested to
Mr. Watson that he should come to Port Erin, obtain

MARINE BIOLOGICAL STATION AT PORT ERIN. 61

living specimens of the worm, and watch its habits in the
Aquarium. Mr. Watson came on August 24th, and the
following day we were fortunately able to take a steamer
to the ground, and amongst the tubes brought up in the
dredge, from over 50 fathoms, one contained a fine living
Panthalis, which was successfully brought on shore and
transferred to a small tank provided with a supply of the
fine mud in which the animal lives. The worm was then
kept under the closest observation by Mr. Watson during
the whole day for the next week, and his enthusiasm and
patient care have been rewarded by the collection of a
number of drawings and notes and some photographs of
the appearance and movements of the living animal.
During that time the Panthalis deserted its old tube and
formed a new one in the mud, fortunately using the glass
for part of one side, so that the process of scooping out,
the mud and of spinning the network of mucus threads
and the various movements of the animal were readily
enough seen—if one did not mind the inconvenience of
lying for hours in a cramped position on the damp con-
erete floor of the Aquarium-room. This satisfactorily
settled the Panthalis question, and established the cor-
rectness of our previous views as to the connection
between the worm and the empty tubes brought up in the
dredge.

Rev. T. S. Lea, M.A., on two visits to the Station,
spent some time in taking photographs of characteristic
specimens of the Marine Algze a sitw at low tide, and in
making microphotographs in the laboratory. He also
photographed some of our collecting grounds and pools,
and some selected pieces of rock showing assemblages of
animals and plants in their natural environment.

We have again been able to send specimens of the
large red mite Bdella bastert to Mr, Michael to aid in his

62 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

work on the Acarina. We have found the home of this
species to be deep in the cracks and fissures of the cliffs
round the Biological Station. From these holes the mites
occasionally emerge in numbers, and are then found over —
the shore and on our concrete steps.

Mr. Chadwick made a special study of the plankton of
Port Erin Bay during part of May, and collected a large
amount of material, which is not yet fully examined. He
found, amongst other things, various stages in the life
history of Amphicodon fritillaria, post larval stages of
Polynoe, Terebella, and Pectinaria, and also the Hal-
stemma-like Siphonophore, which had been found by Mr.
EK. T. Browne in April.

Professor Weiss was at work during the spring at Port
Erin, on Marine Botany. In part he was investigating
the effect of different coloured lights on various living
sea-weeds. He did not arrive at any definite result, but
intends to pursue the matter further. He also spent
some time in examining living Diatoms, with a view to
clearing up some imperfectly known particulars in their
life history. This work is also not yet completed.

Miss lL. R. Thornely and Miss R. Alcock worked
together at the Station for some time in June. Miss
Thornely was occupied with the collection and deter-
mination of Polyzoa, and Miss Alcock was preserving
material for investigations on the microchemistry of the
tests of Tunicata.

In September we obtained some specimens of Calocaris
with ova on the abdomen. The ova are of a pale red
colour, and of relatively large size, measuring about 1°5 to
2mm. in diameter in a specimen about 5 cm. long. There
are from 45 to 60 ova on each individual. It is interesting
to compare these measurements with those of the common
shrimp, where an individual 7 cm. long may have about

}
(

MARINE BIOLOGICAL STATION AT PORT ERIN. 63

5000 ova, measuring each about 0'7 mm. in diameter. So
that in the Calocaris the ova are about three times as large,
but not nearly so numerous, as in the case of its shallow
water relation.

THe ‘‘ Drirt BoTtTLEs.”

In connection with the investigation of the surface life,
in discussing the appearance and disappearance of swarms
of certain Copepods and Medusee, and in considering the
possible influence of the movements of food matters upon
the migrations of fishes, it has occurred to several of us at
different times during the last few years, that 1t would be
worth while to try to ascertain the set of the chief
currents, tidal* or otherwise, such as the movement of
surface waters caused by prevalent winds. The Prince
of Monaco started a few years ago the system of distri-
buting over the North Atlantic large numbers of small
floating copper vessels, with the object of finding out
where they drifted to. This plan we have adopted, with
slight modifications. In September I selected a small,
strong, buoyant bottle, measuring 7°5 cm. by 1°8 cm.,
which seemed well suited for the purpose, and which costs
only 7s. per gross. I drew up a notice, as follows, to go
in the bottles, and had a large number printed and num-
bered consecutively :—

* The tidal currents of the district are already to some extent known, and
ave marked in the charts and given in books of sailing directions, as Admiral
Beechey’s ‘‘ Tidal Streams of the Irish Sea ” but we desire to ascertain the
resultant currents from all influences which would affect the drift of small

floating bodies.

64 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Any one who finds this is earnestly requested
to write the place, and date when found, in the
space (on the other side) for the purpose, place
the paper in an envelope, and post it to

PROFESSOR HERDMAN,
University College,
LIVERPOOL.
Postage need not be prepaid.

(Turn over.

[OTHER SIDE. |
Please write distinctly, and give full particulars.
Hocanrry, where found, ..... .<s:\-+.4:-o060 eee

eeoecerececese eee ree ses eoee eee eee eee oes eee sreeseseeee eee oe essere eeevese Bee

eevee se eee eee ree eee eee eee eee sree eee ese see see eee eee ees eeesese sees eee

Darn, wher JOU 2. 3.20. coe: wodese eee dy
Name and address of sender.

eer eeec oes see ese eee eee eee eee eee sereeeseseeoeer ere eee eee aoe

A paper was then placed in each bottle, so folded that
the number could be readily seen through the glass, the
cork was well pressed down, and dipped in melted paraffin.
Nearly two hundred of these bottles have, since September
30th, been dropped into the sea in various parts of our
area, a record being kept of the locality and time when
each was set free. Several dozen were let off from the
Isle of Man steamer in crossing to Douglas and back, at
intervals of quarter of an hour, and from our trawler when
dredging betweén Port Erin and Ireland. Several dozen
have been let off from Mr. Alfred Holt’s steamers in
going round to Holyhead and in coming down from
Greenock. The fishery steamer “John Fell” has dis-
tributed a number along the coast in the northern part of
the district, and others have been set free at stated

MARINE BIOLOGICAL STATION AT PORT ERIN... 65

intervals during the rise and fall of the tide from the
Morecambe Bay Light Vessel, and from the Liverpool
North-west Light Vessel. Altogether, over 30 per cent.,
or nearly one in three of the bottles distributed have been
subsequently picked up on the shore and returned to me.
They come from various parts of the coast of the Irish
Sea—Scotland, England, .Wales, Isle of Man, and Ireland.
Some of the bottles have gone quite a short distance,
having evidently been taken straight ashore by the rising
tide. Others have been carried an unexpected length,
e.g., one (No. 35), set free near the Crosby Light Vessel,
off Liverpool, at 12-30 p.m., on October 1st, was picked
up at Saltcoats in Ayrshire, on November 7th, having
travelled a distance of at least 180 miles* in 37 days;
another (H. 20) was set free near the Skerries, Anglesey, on
October 6th, and was picked up one mile N. of Ardrossan,
on November 7th, having travelled 150 miles in 31 days;
and bottle No. 1, set free at the Liverpool Bar, on Sep-
tember 30th, was picked up at Shiskin, Arran, about 165
miles off, on November 12th. On the other hand, a bottle
(J. F. 34), set free on November 7th, at the Ribble
Estuary, was picked up on November 12th at St. Anne’s,
having gone only 4 miles.

It would be premature as yet—until many more dozens or
hundreds have been distributed and returned—to draw any
conclusions. It is only by the evidence of large numbers
that the vitiating effect of exceptional circumstances, such
as an unusual gale, can be eliminated. Prevailing winds,
on the other hand, such as would usually affect the drift
of surface organisms are amongst the normally acting
causes which we are trying to ascertain. I may state,
however, that so far nearly 50 per cent. of the bottles

* More probably, very much further, as during that time it would
certainly be carried backwards and forwards by the tide,

66 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

found had been carried to Ireland, and they are chiefly
ones that had been set free in the southern part of the
district (between Liverpool and Holyhead) and off the
Isle of Man. The bottles set free along the Lancashire
Coast and in Morecambe Bay seem chiefly to have been
carried to the south and west, e.g., to about Mostyn, in
North Wales, and Douglas, Isle of Man. It is only a few
exceptional ones so far that have been carried out of our
area through the North Channel.

Plate I. shows by the dark dashes the lines along which
the drift bottles have been set free, and the arrow heads
give some idea of the direction which some of the bottles
have apparently taken.

PUBLICATIONS.

Sufficient material has now been accumulated to form
a new volume of the “ Fauna of Liverpool Bay.’~- Con-
sequently, Volume IV. will be issued early in 1895, and
will contain,.1n addition to the reports and papers which
were mentioned last year as printed off, the Supplemen-
tary Report on the Hydroid Zoophytes, by Miss L. R.
Thornely ; a list of the Nemertida found at Port Erin, by
Messrs. Beaumont and Vanstone; a Revision of the
Nomenclature and Classification of British Sponges, by
Dr. Hanitsch ; a Report on the Fishes of the District, by
Professor Herdman; a paper on Synapta, by Mr. Chad-
wick, and one on the Tube-building Habits of Panthalis,
by Mr. Watson; a report on the L. M. B. C. Meduse, by
Mr. Edw. T. Browne; and a Supplementary Report on
Copepoda, by Mr. I. C. Thompson.

It has been suggested that a general index should now
be drawn up to the species recorded in the volumes on
the Fauna and Flora of Liverpool Bay. There can be
no doubt that such an index would be useful, and will

MARINE BIOLOGICAL STATION AT PORT ERIN. 67

eventually be necessary, but as the work of identifying
species is still in active progress, and fresh records are
being made annually, it 1s perhaps a little premature to
undertake such a piece of work now. Probably an appro-
priate time for the compilation of such an index would be
at the conclusion of the labours of our British Association
Committee, which will probably submit its final report on
the occasion of the visit of the British Association to
Liverpool in the autumn of 1896.

The surplus copies of the five Annual Reports upon the
Marine Biological Station formerly on Puffin Island (1888
to 1892, the complete set) have been collated and bound
up to form an 8vo. volume of about 180 pages, illustrated
with cuts and plates, and containing the original litho-
eraphed covers. There are 20 copies of this vol. which
are now offered by the Committee at 3s. each nett (post
free); apply to Mr. I. C. Thompson, 4, Lord Street, or to
Prof. Herdman, University College.

EFutTURE WORK.

During the coming year the Committee propose, in the
course of their dredging, trawling, and tow-netting inves-
tigations, to pay special attention to the nature of the sea-
bottom and to the assemblages of animals and plants
found on particular deposits. They hope, with the
co-operation of Mr. Clement Reid, and of any local
Geologists who will help them, to work up more
thoroughly the Geology of the sea-bottom, and so attempt
the solution of that interesting problem, the date and
circumstances of formation of the Irish Sea and of the
separation of the Isle of Man from England. ‘They pro-
pose also to continue for at least a year the observations

bd

on currents by means of ‘‘ drift bottles,’ and would be

glad of any assistance in this from shipowners and captains,

68 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

especially in distributing bottles along lines crossing the
northern and southern entrances to the Irish Sea, such as
from Port Patrick to Belfast, or from Belfast to White-
haven, and from Holyhead to Dublin, or thereabouts.

A further subject upon which the Committee are very —
desirous of doing some work is sea-fish hatching and fish
culture. That, however, is a somewhat expensive matter,
requiring special arrangements and apparatus. If the
various County Councils and Fishery Boards having
jurisdiction over the territorial waters of the Irish Sea,
could see their way to unite in promoting a Fish Hatchery
at Port Erin, it would be a benefit to the whole sea-
district. This northern area of the Irish Sea, in the
centre of which the Isle of Man lies, is a natural fishery
district, with its own spawning grounds, “ nurseries,”
feeding grounds, and important fishing industries.. The
same bye-laws ought to apply throughout the area, and
there ought to be a central fish hatching and rearing
establishment in some convenient spot where the water
is the purest obtainable and as free as possible from all
suspended matters. Such a hatchery would not be for
the benefit of Lancashire alone, nor of Cheshire, nor of
the Isle of Man, but would benefit the fish population of
the entire area, and the fisheries in general. TheL.M.B.C.,
although very willing to advise, to co-operate, and to super-
vise if desired, cannot be expected unaided to carry out
such an extensive scheme; but even if they receive no
outside assistance, they are determined to start the matter
by conducting some experiments in hatching during the
coming spawning season, so as to demonstrate the suit-
ability or otherwise of Port Erin for this purpose. In this
connection the Hon. Treasurer begs to call attention +6:
the fact that his small balance in hand will very soon be
exhausted in making the necessary preparations for these

MARINE BIOLOGICAL STATION AT PORT ERIN. 69

experiments, and that he would be very glad to receive
subscriptions for this or any other of the special purposes
mentioned above.

The usual statement of the Hon. Treasurer, and the
the list of subscriptions and donations for 1894 is appended
to this Report.

wT. nite
riya 11,4 Be,
c tN NG S294

SNe Ihe SF
Ni Py
ea W0i5 —
er 5 144 ag a.

D SN ing, A
< wos 44,4 Pig ine y
NS ee th I Wh 4 iat mit, ¢
ee Pee VN NN 089 !
tt NN NN anes )

Cx, 1 \ \ pbnve .
Artie + tH VIN Ce) Ges
wen. | gf a3
Sh at Ne PNY Fy 1 m7
kB: se GA SY, Ww)
< - We Ss ar? 4G
CS SATS, g

An abnormal specimen of Porania pulvillus, where one of the rays shows a
tendency to branch dichotomously. 4, B, C, D are normal rays, while
the fifth ray has divided into x and y, the latter of which has split and
partially united at least twice. The specimen is in the Zoological
Museum of University College, Liverpool. (For the use of this block
we are indebted to the courtesy of Messrs. Macmillan & Co.)

70

APPENDIX.

SUBSCRIPTIONS and DONATIONS.

Subscriptions. Donations.
£ s. 0d.) agen
Alcock, Miss, Cambridge ... . Te =

Banks, Prof. W. Mitchell, 28, aa 2 2h —
Beaumont, W. I., Cambridge oa See —
Bickersteth, Dr., 2, Rodney-street... RS I =
Boulnois, H. P,, 7, Devonshire-road, Prince’s

iPatk 2 “ ae i. ==
Brown, Prof. J. eaten University

College, Liverpool... He .. lo 1
Browne, Edward T., B.A., 14, Uxbridge .

road, Shepherd’s Bush, London 3 —
Caine, Nath., 10, Orange-court, Castle-street 1 1 0 —_
Caton, Dr., 31, Rodney-street .... .o = I gdsbat

Clague, Dr., Castletown, Isle of Man Ls —
Clague, Thomas, Bellevue Hotel, Port Erin 1 1 0 —*
Comber, Thomas, J.P., Leighton, Parkgate i 1 0 as
Crellin, John C., J.P., Ballachurry, Andreas,

Isle of ee ee ; ere ee) —
Dawkins, Professor W. Boel Oph Goines

Manchester tLe —
Denny, Prof., Firth Dales. ‘Sheffield I< loaee —
Derby, Karl of, Knowsley .. aoe ; 8 0a -—
Gair, H. W. ig Sea Wwe 2 2 o =

eres Col David, C.B., Windleburst
St. Helens

Gamble, F. W., Owens Ualeed Mancubies

Gaskell, Frank, Woolton Wood,

Gaskell, Holbrook, J.P., Woolton Wood

Gell, James 8., High Bailiff of Castletown...

ls Se SS)
- = SS ©
So ore S&S

MARINE BIOLOGICAL STATION AT PORT ERIN. 71

Gibbons, Fredk., 19, Ranelagh-street l
Gibson, Prof, R. J. H.,41,Sydenham-avenue 1
Gifford, J., Whitehouse terrace, Edinburgh 1
Glynn, Dr., 62, Rodney-street ee
Greening, Linnzus, 5, Wilson Patten-street,

- Oo =
Sag =>
|

Warrington lapel. 0 —
Hall, Walter J., Lindum House ee 010 0 ==
Halls, W. J, 35, Lord-street Deke 6 ss
Henderson, W. G., Liverpool Union Bank 2) +0 a
Herdman, Prof., i College, L’pool. 2 2 0 —
Hick, Herbert E., M.R.C.S., Southport

House, Bradford Bs : OF 16, 6 =
Hick, Thomas, B.A. LBeeheecare Bas

holme, Ri bticater oe OOS —-
Holder, Thos,, 1, Clarendon pears ithe.

. ee dts big, O —
Holland, Walter, Mossley Hill. te yet ok eo —
Holt, George, J.P. Sudley, Mossley Hill ... 1 0 0O —
Howes, Prof. G. B., Royal College of

Science, South Kensington, London... 1 1 0 —
Hoyle, W. EH., Museum, Owens College,

Pnclicater wi lis skh) —
Isle of Man Natural History and renee

ian Society Et. —
Jones, C. W., J.P., Field Hoge eee 1, Oi. 0 —
Kermode, p M, C, Hill-side, Ramsey. Pele 0) =
Lea, Rev. TT’. Simcox, 3, Wellington-fields 1 1 0 —
Leicester, Alfred, 30, Weld-rd., Birkdale... 1 1 0 a
Lomas, J., Amery-grove, Birkenhead es —
Macfie, abort, Airds 1o7-0:, 0 —
Meyer, Dr. Kuno, University Salles iy eel 0 5 0 —-
Meade-King, H. W., J.P., Sandfied Park 1 1 0 —
Meade-King, R. R., 4, Oldhall-street 0:10. .0 —
Melly, W. K., 90, Chatham-street... ie 2 —
Miall, Prof., Yorkshire College, Leeds ee eee _
Monks, F. W., Brooklands, Warrington et 6 —

72 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Muspratt, E. K., Seaforth Hall

Newton, John, M.R.C.S., 44, Rodney-street
Odgers, Rev. J. Hi., Woodstock Rd., Oxford
Poole, Sir James, Tower Buildings
Rathbone, 8. G., Croxteth-drive, Sefton-park
Rathbone, Mrs. Theo., Backwood, Neston
Rathbone, Miss May, Backwood, Neston ...
Rathbone, W., M.P., Greenbank, Allerton
Reading, N. C., Wake Green-road, Moseley
Roberts, Isaac, F.R.S., Tunbridge-wells ...

Shaw, Prof. H. 8. Hele, 26, Waverley-road

Shepheard, T., Kingsley Lodge, Chester ...
Simpson, J. Fone Annandale, Aigburth-
drive: a.
Smith, A. T., junr., 24, eee!
Stevenson, W.A., Ballakreighan, Castletown,
Isle of Man
Talbot, Rev. T. U., 4, Ota terrace, Dane
glas, Isle of ee 1 a se
Thompson, Isaac C., 19, Waverley-road
Thornely, James, Baycliff, Woolton
Thornely, The Misses, Baycliff, Woolton ...
Toll, J. M., 340, Walton Breck-road
Tomlin, B., 59, Liverpool-road, Chester...
Walker, Alfred O., Nant-y-glyn, Colwyn Bay
Walker, Horace, South Lodge, Princes-park
Walters, Rev. Frank, B.A., King William
College, Isle of Man
Watson, A. T., Tapton-crescent, Sheffield
Weiss, Prof. F. K., Owen’s College, Man-
chester
Westminster, Duke of, Baton Hall
Wiglesworth, Dr., Rainhill...

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74 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

i M.B.C; INOMICEs:

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Aquarium from 12 to 1 and from 6 to 6-30 p.m. daily,
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The L.M.B. Committee are publishing their Reports
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there have appeared :—

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MARINE BIOLOGICAL STATION AT PORT ERIN. Ga

EXPLANATION OF THE PLATES.

Puate I.—Chart of the Irish Sea showing the zones of
depth (contours 10, 20, and 50 fathoms), the nature of the
bottom (see p. 287), and the lines of distribution (thick
dotting) of the drift bottles.

PuatE IJ.—Calcareous “‘concretion,”’ containing shells,
sand grains, &c., cemented with carbonate of lime and a
little carbonate of iron (see last report, p. 31). From a
photograph by Rev. T. 8. Lea.

76

DESCRIPTION of

THREE SPECIES of ANGUILLULIDA, observed

in DISEASED PSHUDO-BULBS of TROPICAG
ORCHIDS.

By Dr. J. -G. de Man;

Or IERSEKE, ZEELAND, NETHERLANDS.

Plates I11—V.

In the beginning of February, 1894, Dr. J. Ritzema
Bos, of Wageningen, Netherlands, sent me some diseased
pseudo-bulbs of tropical orchids, belonging to the genus
Calanthe, R. Br., begging me to inquire, together with
him, into the cause of the disease. We did not succeed
in discovering it, though it became very likely that some
fungus which propagates in the tissue of the said pseudo-
bulbs ought to be considered as the origin of the pest.
In the disorganized tissue, however, which in some
_pseudo-bulbs was changed into a humus-like substance,
several species of Nematode worms were observed: three
of the latter will be described by me in the following
pages.

The pseudo-bulbs had been sent to Mr. Ritzema Bos
from the greenhouses of the Duke of Westminster, at
Chester, England. Some belonged to Cal. vestita, Wall. ;
most of them, however, to Cal. Vertchu, a hybrid form
between the said Cal. vestita and Cal. rosea. The species
of this genus of plants are found in India and Australia,
so, e.g., Cal. vestitta in Burma. I therefore at first~
supposed that the original habitat of these worms was
not Europe, but that they should prove to have been
imported, in the tissue of the plants, from India, especially

ANGUILLULIDAD FROM TROPICAL ORCHIDS. 717

since two of the three observed species are new to science,
and because the third is very likely identical with a
species that inhabits the Fiji Islands.

Mr. R. Newstead, Curator of the Grosvenor Museum
of Natural History, at Chester, however, wrote the
following to Mr. Ritzema Bos :—

“There is absolutely no means of tracing the original
habitat of the Calanthes, for they have been in the col-
lection at Eaton for a number of years. Originally, some
years ago, there were comparatively few plants, but the
orchid grower, Mr. C. E. Hann, has annually increased
the stock by the propagation of the pseudo-bulbs, until
last year, when the collection consisted of about 1,000
plants, of which there is not a single healthy specimen
left. The soil, or ‘potting material,’ used in the cultiva-
tion of the plants, consisted chiefly of turfy-loam, with a
little well-rotted cow dung. When the plants were in
active growth, they were supplied with both artificial and
ordinary liquid manure; the latter was obtained from the
immediate neighbourhood, as also was the soil.”

We ought, therefore, to conclude, as well from these
words as from the fact that Cal. Veitchii, to which most
of the diseased pseudo-bulbs belong, is a hybrid form,
cultivated in the nursery of Veitch, at Chelsea, London,
that my first supposition of the importation of these
worms out of the tropics 1s highly improbable.

1.—APHELENCHUS TENUICAUDATUS, n. sp.—Fig. 1.

Male 0°8 mm., female 0°95 mm.; a = 85—36; P in the
male 83}—9, in the female 9—9}; y* in the male 11—15,
in the female 73—84, rarely 10.

* a indicates the proportion between the body-length and the average
diameter of the body; 8 the proportion between the body-length and
the length of the cesophagus, the pharynx included ; y finally the proportion
between the body-length and the length of the tail,

78 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Both the male and the female of this new species have
a moderately slender body. The body tapers slightly
anteriorly, so that the diameter of the head at the con-
stricted neck measures one-third of the greatest width of
the body. The tail has a very characteristic shape. That
of the female (Pl. III, fig. 1 d) almost constantly a little
longer than the cesophagus, appears slender, elongated,
tapering regularly with a hawr-like termination. The tail
of the male (Pl. IV, fig. 1 c) is constantly shghtly shorter
than the cesophagus, and tapers rather gradually along
its anterior two-thirds, behind the papille suddenly much
more, the termination fine, though not so hair-like as in
the female. There is no caudal gland. The cuticula, as
is usually the case with the species of this genus,
destituted of set, is very finely striated; it 1s very thin,
measuring only 0°9—1 », and the striz are so fine that it
was only at the anterior extremity of the body, opposite
the spear, that I succeeded in observing them distinctly.
A lateral membrane (‘‘Seitenmembran”’) is probably
wanting. |

The truncated head (Pl. III, fig. 1 6) presents six
rounded lips or lobes at the periphery, that may best be
studied in a front view of the head. These lips bear no
papille. The spear is 24—27 » long, cylindroid, without
bulbs at its posterior extremity, and obliquely cut off at
the sharp anterior end like the spear of the species of the
genus Dorylavmus. The wall that surrounds the spear
and that is a continuation of the muscular wall of the
cesophagus is somewhat thicker than the latter. I believe
I have observed a spear-collar, as is generally found in the
species of the genus Dorylaimus. ‘Two long muscles,
that originate from the body-wall near the neck-like
constriction of the head, are attached at the posterior
extremity of the spear, and serve evidently to protrude

ANGUILLULIDA FROM TROPICAL ORCHIDS. 79

it. The conspicuous chitinous tube of the cesophagus, as
usual a direct continuation of the spear, is cylindroid,
the transverse optical section appearing circular, and the
wall of this part of the cesophagus is rather thin. The
latter terminates in a rather large bulb, which 1s one
and-a-half as long as it ts broad, and which measures
almost one-fourth of the distance between its posterior
margin and the mouth; the bulb-wall is distinctly mus-
cular, and the dilatation of the central tube is 5 » long
and 3,6 broad. Behind the bulb the alimentary canal
continues, at first very narrow, but afterwards gradually
widening. The wall of the intestine contains mostly
numerous fat-granules. The excretory pore lies usually
just behind the bulb, sometimes, however, opposite its
posterior portion (fig. 1 a).

I observed (Pl. III, fig. 1) next to the anterior part of
the intestine, on its dorsal side, a gland of an elongated
form, the nature and the function of which, however,
remain unknown. ‘This gland is considerably longer than
the oesophagus; in the female nearly one and-a-half as
long, and includes a circular nucleus, which is 9 » broad.
A more careful examination is necessary to elucidate the
signification of this organ. ‘The genital tube of the male
is simple, and sometimes it seemed to me that the blind
end was reflexed for a short part. The rather large
spicula (Pl. LV, fig. 1 c) have a characteristic form. Their
length, 2.e., the linear distance between their two extremi-
ties, measures 18; they are slightly arcuate, the proximal
ends are capitate, and the distal ends are obliquely cut
off, whereas they bear a triangular prominence on their
anterior margin. An accessory piece 1s wanting. I
observed a little in front of the anus, a single conical
papilla, situated in the ventral middle hne of the body,
and the tail itself bears two pairs of sub-ventral papillee ;

80 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

these two pairs are lying immediately behind one another,
somewhat posterior to the middle of the tail, which, as I ~
have already remarked, tapers considerably behind the
papille, to terminate in an acute point. The vulva is
situated rather far backwards, and its distance from the
extremity of the tail measures constantly a little more
than one-fourth and a little less than one-third of the
whole body-length, whereas the distance between the vulva
and the anus appears always nearly one and-a-half times
as long as the esophagus and longer than the tail. The
thin-walled vagina extends obliquely forward. The sexual
tube, that is not reflexed, extends till near the anterior
end of the intestine, and the uterus bears a posterior
branch, which is a trifle longer than half the distance
between the vulva and the anus (PI. Ul, fig. 1). This
species is oviparous. The eggs are slightly arcuate, a little
more than three times as long as broad, and 0°07—0:075 |
mm. long; they are rounded on the extremities.
Numerous individuals, both males and females, as well
as young asexual larve, were observed by me in the
dis-organized humus-like tissue of the Calanthe. The
movements of these worms are rather sluggish. |
From all other species of the genus A phelenchus hitherto
observed in Europe, Aphel. tenwicaudatus at first sight
may be distinguished by the elongated, slender, finely
ternunating tail. Last year, however, another new
species of this genus was described by Mr. Cobb,* under
the name of Aph. longicaudatus, in which the tail-end
is likewise hair-like in both sexes, and appears even com-
paratively somewhat longer than in Aph. tenuicaudatus.
In that species discovered by Cobb in soil about banana
* Cobb, Nematodes, mostly Australian and Fijian, Sydney, 1893, p. 54.,

Pl. VII. (Department of Agriculture, New South Wales. Miscellaneous
Publications, No. 13.)

ANGUILLULIDA FROM TROPICAL ORCHIDS. 81

plants, on the Fiji Islands, the tail attains nearly one-third
of the whole body-length. It is thus considerably longer;
but, moveover, Aphel. longicaudatus differs in the situa-
tion of the vulva, the distance between the vulva and the
anus being not longer, but only half as long as the tatl.

2.—RHABDITIS CORONATA, Cobb. Fig. 2.

Rhabditis coronata, Cobb, Nematodes, mostly Australian
ae qian Sydney, 1893, p. 30, Pl. ITI., fig. 1.

Male 0°5 mm., female 0°6 mm.; @ in the male = 15,
in the female = 16—17; 8 in the male 43, in the female
48; yin the male = 23, in the female = 74—8.

I identify this species, which differs from all the other
numerous representatives of the genus in the singular
structure of the head, with Khabd. coronata, Cobb, though
not with full certainty, because the description as well as
the figures published by that author, leave much to desire
as regards exactness and completeness. The principal
character is described by Cobb in the following words :—
“The head was surmounted by six (?) conical lips, each
turned outwards,’ and for the rest, the dimensions also
agree rather well, when we consider that only a young
individual was observed by him. The single specimen, a
female, observed by the Sydney naturalist (the male
remaining unknown) had only a length of 0°36 mm.

The body of these worms 1s of a rather stout and plump
shape, and tapers at first only a little anteriorly, but from
the middle of the cesophagus towards the head (Pl. III,
fig. 2) with increasing rapidity, so that the diameter of the
head at the base of the lips measures only one-fourth of the
body-diameter at the commencement of the intestine.
Opposite the anus the females are just half as broad as at the
posterior extremity of the cesophagus, but the males are
somewhat broader at the anus, as usual. ‘lhe cuticula is

82 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

striated, but so very finely, that the strize could be observed
by me with certainty, only at the head. The lateral
membrane (Seitenmembran), however, is very conspicuous
and broad, as it 1s 5°8 » broad in the middle of the body,
v.e., one-sixth of the body-diameter.

As I have already remarked, this species is distinguished
from all others by the structure of the head. I did not
succeed, however, in elucidating that structure thoroughly |
and completely. The head is not formed by six lips, as
was supposed by Cobb, who very likely has studied his
single specimen only in the lateral position, but by four,
namely, by two pairs of lips, which have a very different
structure. One lip is situated in the dorsal median line,
the other lp of the same pair stands, opposite the
former, in the ventral median line. The anterior or oral
surface of these two lips, which is non-transparent and
appears rather dark, is slightly convex, and terminates
externally with two sharp teeth, which are separated from
one another by a concave emargination; these teeth, of
course, become at once visible in a front view of the
head. The posterior wall of these lips, by which they
are separated from the neck, 1s also thickened, and appears
consequently likewise dark, but the concave external wall
is not darkened (Pl. IV, fig. 2a). On account of their
outer wall being concave, both lips appear in the usual —
lateral position, as ‘“‘conical lips turned outwards,” as
writes Cobb, and in that position they are at once
conspicuous, even when only a little magnified, in con-
sequence of the dark colour of their anterior and posterior
walls.

The shape and the structure of the two lips of the
other pair that stand laterally, did not become fully clear
to me, which ought to be ascribed to the fact that these
lips are not chitinous nor thickened, and consequently

ANGUILLULIDA FROM TROPICAL ORCHIDS. 83

transparent, so that it remained impossible to distinguish
distinctly their contours. These lateral lips (Pl. III, fig.
2 b) are considerably Iigher than the lips of the dorso-
ventral pair, and project, therefore, slightly beyond them ;
their outer wall is slightly convex, and their anterior
margin seemed to be concavely emarginated. The reader
will do well to consult the drawings on Plates III and IV.

The mouth leads into the pharynx, which seemed to be
eylindroid ; it is 22 » long in the male, 24—26 p» long in
the female, and the two lines of its longitudinal optical
section run parallel. The pharynx appears nearly eight
tumes as long as it 1s broad, including the wall. According
to Cobb, the pharynx should be ten times as long as it is
broad, but I presume that he did not include the wall.
The length of the pharynx measures thus nearly one-fifth
of the distance between the posterior extremity of the
oesophagus and the mouth. The cesophagus (Pl. III,
fig. 2) has the same form as in most other species of
Rhabditis. The foremost part, between the pharynx and

the median bulb, is 29—30 » long, slightly longer than

the pharynx, and nearly three times as long as it 1s wide.
The next part, the ellipsoidal median bulb, is little more
than half as long and about as wide asitislong. ‘The fol-
lowing portion, between the two bulbs, has about the same
length as the pharynx, but it 1s nearly twice as wide;
this portion is also half as wide as the median bulb. 'The
posterior bulb is 22 » long, and spherical ; it is accordingly
a trifle shorter than the pharynx, and does not yet
measure one-fifth of the whole length of the cesophagus,
the pharynx included. The wall of the cesophagus con-
tains the usual muscular fibres, and the chitinous valvular
apparatus is well developed. The ventral excretory pore
is situated opposite the latter.

Though very many female individuals were found in the

84 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

humus-like substance of the diseased Calanthe, only one
single male specimen was observed by me. The tail (Pl. V.
fig. 2 d) is not completely enveloped by the bursa, but its
extremity seemed to be broken off, so that the terminal
part perhaps afterwards will prove to be longer. The
exact number of the ribs or papillae remained also uncer-
tain. Immediately posterior to the anus lies a rib, that
seems to reach the margin of the bursa, when the latter
is looked at in the usual lateral position; the second and
third rib, the latter of which seemed to reach the margin,
the former not, are situated close together at some distance
from the first. The fourth rib, which reaches the margin,
hes as far distant from the third as the third from the
first; and, finally, two or three ribs were still observed
near the posterior end of the bursa. The bursa is thus
doubtless supported by at least six or seven ribs, but it
remained uncertain whether there are also preanal ribs.
As I have already observed, the body is a little broader
opposite the anus than in the female. The slightly
arcuate spicula are 27 » long, their proximal ends are ©
capitate, and they seemed to narrow to an acute point. —
The lamelliform accessory piece is not yet half as long as 3
the spicula, measuring 11». The blind end of the genital
tube is reflexed: the distance between the posterior end
of the cesophagus and the flexure measures about one-
third of the length of the intestine.

The vulva is constantly situated slightly posterior to
the middle of the body. The two branches of the sexual —
apparatus, on either side of the vagina, have about the

same length, and extend respectively along two-thirds of

the distance between the vulva and the posterior bulb of
the oesophagus, and of that between the vulva and the ~
anus.

Innumerable eggs were observed lying free in the

ANGUILLULIDEH FROM TROPICAL ORCHIDS. 85

4 _ humus-like substance; they are 43—51 » long and about
| half as broad (ig. 2). The tail of the female (Pl. IV,
fig 2 e) is slender, elongated, and tapers regularly towards
the hair-fine termination ; 1t shows the peculiar character
that at a shght distance behind the anus the internal layers
of the cuticula diverge from the external, being separated
from one another by a light coloured one.

The movements of these graceful worms are about as
active as those of the other species of this genus.

Ethabditis coronata was discovered by Cobb in humus
about the roots of banana plants, Fiji Islands, July, 1891.

3.—RHABDITIS OXYCERCA, n. sp.—Fig. 3.

Male and female 0°97 mm.; ain the male 16—1734, in
the female 15—21; 8 = 4—44; yin the male 23—27, in
the female 35—45.

This interesting new species, numerous individuals of
which have been observed by me, both males and females,

| in the diseased tissue of the Calanthe, presents (especially

on account of the shape of the tail in the female) some
resemblance with Rhabd. teres, Schn., but is distinguished
from all other species by the quite different distribution
of wits eight bursa-ribs, and, moreover, by some other
characters.

Lthabditis oxycerca, thus named after the short mucro-
nated tail m both sexes, attains scarcely the length of one
millimeter, and ought accordingly to be reckoned among
the species of small size. The body of these parasites
appears more or less stout, plump, not slender, both in
the male and in the female, though the general form of
the body of the female varies somewhat according to the
uterus being filled or not with eggs and embryos. The
anterior cesophageal part of the body tapers rather much
anteriorly, so that the head-diameter at the base of the

86 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

lips measures only one-fourth of the width of the body at the
commencement of the intestine. Posteriorly the body
tapers only slightly to the anus, and in the male even in a
less degree than in the female.

The cuticula is moderately thick, measuring 2—3 » in
the middle of the body, and is formed by several layers.
Externally, three very thin layers are found, which are
followed by a thicker one, and the latter finally is still —
separated internally from the muscular layer by one or
two that are also very thin. The cuticula is smooth, not
annulated, at least the outermost layers, but the internal
thin layers are, perhaps, very finely striated. The
described thick middle layer of the cuticula is still con-
siderably more thickened on the tail; in the male it
produces the bursa, and in the female the singular form
of the tail is occasioned by it. When these worms, while
swimming, bend and curve their bodies, transverse plaits
are formed by the outermost layers of the cuticula, similar
to those that were described by Claus for Rhabd. brevi-
spina, In the male a very narrow lateral membrane was
observed, only 5 » broad in the middle of the body, 2.e.,
one-twelfth of the body-diameter ; in the female I did not.
observe it positively, but I think the female will also
prove to be provided with it. The rather broad lateral
fields measure one-third of the body-diameter. ,

The head of our species (PI. V, fig. 3a) is surmounted

by six lips, that are separated from one another by rather
shallow notches. The lips are only 3°6 » high, this height
being about equal to one-fourth the diameter of the head at
the base of the lips; the latter are accordingly low, and each

is surmounted by an extremely small papilla, that becomes —

only conspicuous when highly magnified (Seibert, Immer-
sions-system, No. VIII). The lips are separated by a- q
hardly perceptible constriction from the contiguous part

ANGUILLULIDA FROM TROPICAL ORCHIDS. 87

of the neck. The very small, circular lateral organs are
situated immediately behind the lip-region; their diameter
is only 15. The chitinous pharynx is 25°5—27 p» long
from the mouth to its posterior end, and this length
measures from one-ninth to one-eighth of the distance
between the mouth and the commencement of the intes-
tine. The two chitinous lines, which ave the longitudinal
optical section of the pharynx-wall, run parallel; the
pharynx is wide, so that its transverse diameter measures
one-fifth of its length. About in the middle it is sur-
rounded by a very thin, circular, chitinous band, and at
its posterior extremity it presents the ordinary constriction.

The cesophagus (Pl. IV, fig. 3) has the same form as in
many other species of this genus, as e.g., in Rhabd. brevi-
spina, Biitschlii, curvicaudata, &c. The foremost portion,
which has a moderate width, leads about in the middle to
the ellipsoidal median bulb, which afterwards gradually
narrows to terminate with the sharply separated off
posterior bulb, which is nearly spherical. The median
bulb is about one and-a-half times as long as it is broad,
and the narrow portion of the cesophagus between the two
bulbs appears nearly six times as long as it is broad.
The posterior bulb, the valvular apparatus of which is
well developed, occupies one-sixth of the whole length of
the cesophagus, the pharynx included. The wall of the
intestine has the usual structure. I will only remark
that the rectum of the female is rather long, measuring
36 », accordingly still somewhat longer than the tail.
The nerve-ring is seen just behind the median bulb.
The ventral excretory pore, situated just behind the
nerve-ring, lies on a papillary prominence of the cuticula;
the excretory duct, when it comes opposite the pore, makes
first a little curve forward, and then, before terminating,
presents a small dilatation (Pl. V, fig. 3 0).

88 TRANSACTIONS LIVERPOOL BIOLOGICAL SOUIETY.

The genital tube of the male, though being exactly as
long as the intestine, does not reach as far as the ceso-
phagus, because one-sixth is refleced ; 1t shows its greatest
width just in the middle, measuring here two-thirds of
the body-diameter, and thence it narrows gradually
towards the anus. The spermatozoa are spherical cor-
puscles, having a diameter of 7:3—8'7 », and enclosing a
nucleus. The spicula are 42 or 48 » long, accordingly a
little longer than the tail; they are entirely separated
from one another, not coalescent, and their proximal ends
are characteristically capitate; they appear slightly arcuate,
and terminate in an acute point. The accessory piece,
12°5 » long, measures scarcely one-third of the length of
the spicula; it is swleate or horse-shoe like, the opening of
which is directed towards the anus, 2.e., outward, whereas
at the proximal end the two parallel lateral pieces are
united in an arcuate line; these lateral pieces are slightly.
thickened towards their distal extremities. i

Rhabditis oxycerca is chiefly characterised by the
structure of the bursa. In my opinion the bursa of
this species (Pl. IV, fig. 3 ¢ and Pl. V, fig. 3 d) is formed
by the middle layer of the cuticula, described above,
being locally thickened on the tail, consequently not
by lateral wings of the cuticula; this thickened layer
expands also, namely, over the sub-ventral and sub-dorsal
sides of the tail of the male. The tail, nevertheless, is
not completely enveloped by the bursa, but a short,
pointed terminal part projects beyond wt, remaining free ;
this free portion is about half as long as the enveloped
part of the tail. The bursa rs supported by eight papilla,
the distribution of which ws highly characteristic and
exceptional. Six papille are situated on the ventral
surface of the bursa, at some distance from the ventral
median line; these papille are consequently swb-ventral.

ANGUILLULIDZ FROM TROPICAL ORCHIDS. 89

The seventh papilla is situated exactly laterally, the eighth
on the dorsal side of the body, being, therefore, swb-dorsal.
Three of the six pairs of sub-ventral papille are situated
before and three behind the anus. The first and second
of these sub-ventral papillz are placed close together, one
behind the other, near the posterior end of the bursa; the
third is almost as far distant from the second as from the
anus; the fourth, which les immediately in front of the
anus, is slightly more distant from the third than the third
is from the second; the fifth about as far distant from the
fourth as the latter is from the third; but the sixth sub-
ventral papilla is placed considerably further forward, and
is not visible on the figures. The distance indeed between
the sixth papilla and the fifth measures 60 p, that between
the fifth and the first 38 », so that the two distances are
nearly in the proportion of 3:2. ‘The seventh pair of papille
lie exactly laterally, opposite the fourth sub-ventral, a
little before the anus; the last or eighth pair finally is
situated sub-dorsally, on the tail itself, nearly opposite
the second sub-ventral. Whereas, in most other species
of the genus Rhalditis, the bursa-papille are all lateral ;
im Rhabd. oxycerca, on the contrary, most of them are
sub-ventral, being placed on the ventral surface of the
body. It is in consequence of this quite exceptional
distribution of the papille that, in a ventral view of the
bursa, only two of them, viz., the seventh and the eighth,
appear to be situated in the lateral portions of the bursa,
and appear to reach its external margin. In the usual
lateral position of these worms, on the contrary (fig. 8 ¢),
the six sub-ventral papille appear to reach the margin of
the bursa: in this position it becomes also conspicuous
that the bursa does not expand over the whole ventral
surface of the tail, but only over the sub-ventral or lateral
parts of it, so that the ventral surface appears concave,

90 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Opposite the fifth sub-ventral papille, at some distance
in front of the anus, moreover one single papilla is found,
placed exactly in the ventral median line of the body, and
appearing as a very small circular ringlet in a ventral
view of the bursa: a similar median papilla has been
described by Butschii in Rhabd. pellio and Rhabd.
pellrordes.

The vulva, somewhat prominent, like the anus, 1s —
constantly situated at a short distance behind the middle
of the body. The walls of the vagina are rather thick;
nevertheless, they do not present the more or less intense
yellow colour which is often observed in other species,
because they are not chitinous. On each side of the
vagina two rather large, coarsely-granulated, roundish
glands are situated, one before and one behind it, which
empty into the vagina by rather long excretory ducts,
because the glands are lying on the lateral sides of the
body. The sexual organs are double and symmetrically
reflexed. In those adult individuals, in which the embryos
have already left the egg-shells and swim about in the
uterus, the anterior branch of the sexual apparatus extends
along four-fifths of the distance between the vulva and
the commencement of the intestine, the posterior branch,
however, along three-fourths of the distance between
vulva and anus. In those individuals, the uterus of which
contains only eggs, including more or less developed
embryos, the sexual organs extend somewhat less far.
The reflexed ovaries, which soon after the flexure become
narrow, are very long, and extend a little beyond the
vagma.

Rhabditis oxycerca appears to be viviparous. A female
specimen that had lived some time in some humus, taken
from the Calanthe, and mixed with a little water, proved
to contain thirteen embryos, some of which had already

]
fp

ANGUILLULIDZ FROM TROPICAL ORCHIDS. 91.
left the egg-shell, swimming about in the body-cavity of.
the mother, whereas the others were still enveloped by
it ; a few eges were still in various degrees of segmenta-
tion. In the uterus of the other numerous specimens
studied and examined by me, I never observed embryos,
but only eggs in different degrees of segmentation, the
number of which varied from one to nine. These eggs
are 51—55 p long, about half as broad, and both extremi-
ties are broadly rounded.

The tail of the female, resembling that of Rhabd. teres,
Schn., appears bluntly rounded, with conical, pointed ter-
mination, which is about half as long as the distance between
the anus and the extremity of the tail (Pl. V, fig. 3 f).
Similarly, as in that species, the internal layers of the
cuticula diverge from the external, so that it gives the im-
pression that two tails are pushed into one another, and,
just as in Rhabd. teres, on each lateral side of the tail a
minute, distinctly innervated papilla is found, situated in
fthabd. oxycerca, however, just at the base of the free conical
termination. In the body-cavity of most female specimens,
between the muscular layer and the intestine and sexual
organs, a close layer of fat-granules was observed, by
which the said parts are hidden; the alimentary canal
and the sexual organs become, therefore, more or less
inconspicuous in these individuals, so that, é.g., the
valvular apparatus or part of the intestine are often
scarcely visible. I observed these fat granules likewise
in the male individuals, but they surround here chiefly
the posterior half of the cesophagus and the beginning of
the intestine, whereas they are almost wholly absent in
the posterior half of the body, in consequence of which the
portions of intestine and testis which lie there, are much
more conspicuous than in the female.

Also young, just born, asexual individuals were studied.

92 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

When they have a length of 0'4 mm., a is expressed by
19, 8 by 3, and y by 15. The tail of young larvee of this —
size is still conical (Pl. V, fig. 3 h), and terminates in an
acute point. The cuticula is finely striated; the lateral
- membrane very conspicuous, presenting itself as a narrow
line, the width of which is only one fifteenth of the body-
diameter. The lips of the head are already conspicuous,
and the pharynx already almost as long as in the adult
individuals, but it 1s considerably narrower ; the width of
the pharynx, indeed, inclusive of its wall, measures only
one-ninth of its length, so that it appears to be only
nearly half as wide as that of the full developed worms.
The cesophagus presents already the same form as in
the latter ; the median bulb is well developed, as is also
the posterior, the length of which, however, still only
measures one-seventh (instead of one-sixth) of the distance
between the mouth and the extremity of the cesophagus.
I observed already in this specimen the oval rudiment of
the genital organs, which was 13long. In another young
larva, 0°44 mm. long, the said rudiment had already a
length of 18 ». Besides by the striated cuticula, these
larve differ at this size by the pharynx being narrower,
and by the different shape of the tal.

EXPLANATION OF THE PLATES.*

Puate ITI.

Fig. 1. Aphelenchus tenuicaudatus, n. sp., ova-bearing
female, 0°95 mm. long, X the elongated gland, ous la,
anterior cesophageal portion of the body of another female -
individual, 0°9 mm. long, X the ventral excretory pore,

* When the contrary has not been stated, the figures have been drawn in
the usual lateral position of these worms.

- ANGUILLULIDE FROM TROPICAL ORCHIDS. 93

Sos; 1b, head of the same specimen, 13%2; the two

minute points which indicate the spear-collar, are
omitted; 1d, tail of a female specimen, 0°9 mm. long, #35.

Fig. 2. Rhabditis coronata, Cobb, anterior cesophageal
portion of a female specimen, 0°6 mm. long, x the
excretory pore, *2°; the two tinted stripes which indicate
the pharynx Pabht ie be shghtly thinner; 2b, ventral view
of the head of this specimen, 242°.

PuATE LY.

Fig. 1c. Aphelenchus tenuicaudatus, n. sp., tail and
anal region of a male, 0°8 mm. long, showing spicula and
papille, 427°.

Fig. 2a. Ehabditis coronata, Cobb, head of the same
specimen as above, 74°"; 2c, ventral view of the spicula,
200. %e, tail of a ae specimen, 0°556 mm. long, 27°;
of ies ege with embryo, 29°.

Fig. 3. Rhabditis oxycerca, n. sp., anterior cesophageal
500 - in this

figure the lips are not quite symetrically ao and the

portion of the body of a male, 0°38 mm. long,

two stripes which indicate the pharynx wall ought to be
slightly thinner; 3c, tail and bursa of an adult male, +42°.

| PrarTs “V,
Fig. 2d. Khabditis coronata, Cobb, lateral view of the

bursa and of the spicula, 29°.

Fig. 3a. Rhabditis oxycerca, n. sp., head of an adult,
ova-bearing female, the ventral side being on the left
hand, 122°; the transverse line on the middle of the
pharynx ought to be slightly thinner ; 30, terminal portion
of the excretory duct and the pore of a female, +#/°; 3d,
ventral view of the tail and bursa of an adult male, 1870.
3e, ventral view of the extremities of the spicula and of

the distally thickened extremities of the lateral portions

94 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of the accessory piece, 24°°; 3f, tail of an adult female
specimen, X the left lateral papilla, 1370 ; 3g, ventral view
of the terminal portion of the tail of an ova-bearing female,
presenting the two papille, 242°; 3h, tail of a young larva, —
0.44 mm. long, +2£° (the striw of the cuticula are not
drawn). In this figure the margins of the posterior half
of the tail ought to be slightly convex.

95

[WORK FROM THE PORT ERIN BIOLOGICAL STATION. |

RECENT ADDITIONS to the COPEPODA of
LIVERPOOL BAY.

By Isaac C. THompson, F.L.S., F.R.M.S.

With Plates VI and VII.
[Read November 9th, 1894.]

S1IncE the Revised Report on the Copepoda of Liverpool
Bay was published last year, twenty-four species new to the
district have been recorded, one of these, Pseudocyclopia
stephoides, being new to science.

Surface tow-nets have been continuously employed
during the several marine expeditions undertaken by the
Committee, also tow-nets attached to the rope a few
fathoms above the dredge. The latter device has proved
a success, collecting some good species of Copepoda, as
well as Cumacea and Amphipoda, which are seldom or
never obtained on the surface. Amongst the Copepoda
thus obtained were several specimens of Pseudocalanus
armatus, found along with a shoal of Pseudocalanus
elongatus. A widely extending shoal of Anomalocera
patersonu was observed off the Isle of Man in May, the
only occasion on which we have taken this species during
the year. On several occasions, notably in the early part
of June, the surface organisms have been singularly scarce.

Special care has been taken to wash and sieve through
fine silk as much as possible of the material brought up
by the dredge during marine expeditions, and it is by this
means that several of the above-mentioned Copepoda, new
to the district, have been obtained, as well as the new
species Pseudocyclopia stephordes,

96 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Mr. Andrew Scott, ‘‘ Fisheries’ Assistant at University
College, has worked through a large amount of material
collected at low water and on various marine expeditions,
with excellent results, the majority of those here mentioned
having been found by him. To his kindness and skill in
delineation I am also indebted for the drawings from
which the plates accompanying this paper are engraved.

DESCRIPTION OF SPECIES.
COPEPODA.
Family PsEUDOCYCLOPIIDA.

Pseudocyclopia stephoides, n. sp. (Pls. VI and VII,
figs. 1 to 14). |

Length, exclusive of caudal sete, 1:2 mm. Cephalo-
thorax robust, four-jointed, the first segment being two-
thirds the combined length of the other three. Abdomen
five-jointed in the male (fig. 14), four-jointed in the female
(fig. 13); the lower portion of the first joint in the male
abdomen (fig. 14) is covered with fine very short hairs or
spines; the first joint in the female abdomen is about
equal in length to that of the combined succeeding three-
joints. Rostrum short.

Anterior antenne (fig. 2) of moderate length, twenty
jointed. Basal joint large, almost equal in length to the
succeeding six joints, and bearing three plumose setz and
one shorter seta; the seventh, ninth, thirteenth, and
twentieth joints have long sensory filaments; the nine-
teenth joint has one plumose seta; each joint bears one
or more ordinary sete. The proportional lengths of the
joints are about as follows :—

4533456343 3 8 38 38° 3°33
1693456789 10:11 12 13 14 1516 17 tomes

Primary branch of posterior antenne (fig. 3) two-jointed,
in this respect and no other differing from the generic

ADDITIONS TO L.M.B.C. COPEPODA. 97

character of Pseuwdocyclopia, Scott, in which the primary
branch is described as being three-jointed. The basal joint
has two marginal sete, the terminal joint having a number
of apical sete. Secondary branch large, five-jointed, the
first, second, and apical joints being about twice as long
as broad, the third and fourth about half as long as broad.

Mandibles (fig. 4) large, consisting of a broad biting
part furnished with two plumose spines, and a two-
branched palp, one of the branches being two, the other
four-jointed. ,

Anterior foot-jaw (fig. 5) four-jointed, with several
mareinal processes, bearing long sete, some finely plumose,
the third joint having two powerful serrated claw-lke
spines; the fourth joint very small, and terminated with
two long sete. The posterior foot-jaw (fig. 6) 1s seven-
jointed, the basal joint large, about twice as long as broad,
bearing several marginal spines, the upper distal angle
protruding upwards, and terminated by three long sete,
the lower one plumose. The second joint is about equal
in length to the first, and little more than half its width ;
the inner margin clothed with short sete, and having
three long plumose sete; the five terminal joints are
small, their combined length being rather less than the
first or second joints, and all thickly clothed with long
sete.

The swimming feet are very similar to those of P.
crassicorms, Scott. The outer branch of the first pair
(fig. 7) is three-jointed, each joint being provided with a
stout dagger-like spine at the outer distal angle; the inner
branch is one-jointed, and rather longer than the first
joint of the outer branch. The outer branch of the second
pair (fig. 8) is also three-jointed; each of the first and
second joints bear one, and the last joint four, stout
serrated spines of variable length, the terminal one being

98 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

serrated only on the inner side; the inner branch is
two-jointed, about half the length of the outer branch, its
inner joint being about half as long as the outer one. The
densely plumose setz in the second, third, and fourth
swimming feet are all jointed at about one-third of their
length. The third (fig. 9) and fourth pairs (fig. 10) have
both branches three-jointed. The outer branch in both
pairs is very similar to that of the second pair. The inner
branch of the fourth pair (fig. 10) has a strong hairy
spine at the distal angle of the first and second joints in
place of jointed seta in the fourth pair. Hach of the fifth
pair in the female (fig. 12) is one-branched, two-jointed,
the first joint short, about as long as broad, with a spinous
prolongation in the centre on the inner side. A similar
projection, as well as a smaller one occur on each inner
side of the segment from which the fifth feet spring.
The second joint of the fifth pair is produced into three
plumose spines (without articulation), the inner terminal
one being longer than the two outer lateral ones. Hach
of the fifth pair of feet in the male (fig. 11) is one-branched
and four-jointed, and together form a powerful clasping
organ. The right foot is long and slender, the terminal
joint being about the combined length of the other three ;
its centre portion almost forms a semi-circle, the continu-
ation being a long sharp spine. The left foot is shorter.
there are several setze and a short blunt spine on the third
joint; the fourth, which is small, terminating in a curved
claw-like prolongation, with a sharp spine near the apex.

The caudal stylets (fig. 13) in the female are about as
long as broad, those of the male (fig. 14) being rather
longer; each bears three long and one short plumose sete.

Three specimens only, two males and one female, were
found in washings from dredged material taken outside
Port Erin, in 15 fathoms, in March, 1894.

ADDITIONS TO L.M.B.C. COPEPODA. ~~ - 99

It was by no means easy to decide into which genus to
place this well-marked species, as it has strong points of
resemblance in common with the three genera, Pseudo-
calanus, Stephos, and Pseudocyclopia. With Pseudocyclo-
pia it agrees in all points excepting in the number of joints
in the anterior antenne, and the primary branch of the
posterior antenne, and, as in general appearance and in
the first four pairs of swimming feet, it strongly resembles
Pseudocyclopia, I have decided provisionally to place it
in that genus. Its fifth pair of feet, however, are more
like those of Stephos.

In the Twelfth Annual Report of the Fishery Board
for Scotland, Mr. Thomas Scott has added a new species
belonging to this genus, recently found by him in the
Forth area.

As the genus Pseudocyclopia forms a sort of missing
link between the families Calanidz and Misophride,
Scott has wisely constituted a new family, the Pseudocy-
clopiide, for its reception. The species of Pseudocy-
clopia, described by Scott having respectively sixteen
and seventeen joints in the anterior antenne, he has
made that number a family character. ‘The species here
described has, however, twenty joints in the anterior
antenne, and as it otherwise agrees in all respects with
the family characters of Pseudocyclopide, I would suggest
that the words “sixteen to seventeen jointed” be altered
to “sixteen to twenty jointed”’ as a character of this new

family.

Family CYCLOPIDE.

Cyclops magnoctavus, Cragin.

One or two specimens of this brackish species were
found along with quantities of Temorella affinis and
Tachidius brevicornis in tow-net gatherings sent to me

100 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

by My. Ascroft, taken by him in low water marine pools
at Lytham. It is evident that a considerable amount of
fresh-water finds its way into the Lytham pools.

Cyclops ewarti, Brady.

This species, first taken in the Forth estuary, was
suspected by Brady to have a fresh-water origin. Ours
are evidently strictly marine, two specimens, both males,
having been dredged at 20 fathoms off Port Erin.

Family HARPACTICIDA.

Longipedia minor, T. & A. Scott.

A few specimens of this species were collected by hand-
net in the rock-pools at Hilbre Islands in March, by
Mr. A. Scott. It is easily distinguished from L. coronata,
Claus, by its much smaller size.

Canuella perplexa, T. & A. Scott.

Frequently found in dredged material taken about Port
Erin. It has probably been overlooked from its general |
resemblance to Longipedia coronata, the points of differ-
ence being enumerated by Scott.

Ectinosoma normam, T. & A. Scott.

Several specimens were obtained by Mr. A. Scott in
material from Barrow Channel, collected in May by
Professor Herdman, and I have also dredged it off Port
Erin. When fresh this species has a brilliant red spot
on the lower angles of the cephalothorax, and in this
respect it agrees with H. erythrops, Brady.

FEctinosoma elongata, A. & 'T. Scott. .

One specimen was found in material from pools at
Hilbre Island.

Ectinosoma gracile, T. & A. Scott.

One or two specimens of this species were obtained
among dredged material collected at Port Hrin by
- Professor Herdman, 7 |

ADDITIONS TO L.M.B.C. COPEPODA. , TOK

Ectinosoma pygmeum, T. & A. Scott.

This species was obtained from the same material as
the last, and is the smallest known LEctinosoma: it
measures only 7th of an inch (33 mm.).

Ectinosoma herdmant, T. & A. Scott.

One specimen was found in dredged material taken off
Port Erin.

Bradya minor, T. & A. Scott.*

A few species of this new Bradya were obtained in
rock-pools at Hilbre Island, along with Longipedia nunor.

Amevra longicaudata, Scott.

One specimen was found in material dredged at 15
fathoms between Port Erin and Peel.

Dactylopus rostratus, 'T. Scott.

A single specimen was obtained among some dredged
material collected at Port Erin by Professor Herdman, at
Faster, 1894.

Diosaccus propinguus, T. & A. Scott, Ameira exigua,
T. Scott, A. longirenis, T. Scott, Laophonte inopinata,
1’. Scott, Pseudowestwoodia pygmea, T. & A. §., and
possibly a new Laophonte, and one or two other doubtful
species were obtained from washings from sponges col-
lected by Dr. Hanitsch at Port Erin in August, 1894.

Family HERSILIIDA.

Cancerilla tubulata, Dalyell.

The first record of this rare Copepod occurs in Dalyell’s
“Powers of the Creator,’ 1851, and it has since been
taken by Mr. Gamble at Plymouth, but not before in our
district. I found it lately on examining the results of

* The above species of Hetinosoma and Bradya are figured and described
in a revision of the British species of Copepoda belonging to the two genera

Letinosoma and Bradya, T. & A. Scott, which is to be published at an early
date.

102 TRANSACTIONS LIVERPOOL BIULOGICAL SOCIETY.

a recent expedition from ‘Port Erin. Large quantities of
ophiuroids, chiefly Ophiocoma nigra and Ophiothria
fragilis, were amongst the dredged material, and it is
probably from one or other of these that the two speci-
mens of Cancerilla tubulata, Dalyell, male and female,
were taken, as the species 1s parasitic on ophiuroids. It
has recently been fully described and figured in “‘ Les
Copépodes du Boulonnais,”’ by Dr. Kugene Canu.

Family SAPPHIRINIDZ.

Pseudanthessius sauvager, Canu.

A few specimens were obtained by washing a number
of Spatangus purpureus, which were trawled in the
central area, 21 miles W.N.W. from Morecambe Bay
Lightship, on April 3rd. This rare species was only
added to the British fauna last year, when it was found
in the Firth of Forth, and the present is the second time —
it has been observed in the British area.

Lichomolgus (Doridicola) agilis, Leydig, was found in
the bottom tow-net, Morecambe Bay, May, 1894.

Family ARTOTROGIDA..

Acontiophorus elongatus, Scott. —
One specimen was found among the strained washings
of Pecten maximus, dredged at 15 fathoms, off Port Erin.

Family CALIGIDA.

“essere Po rons: :
Several specimens, male and female, were found on ihe
flounder, taken off Morecambe, and also from Arnoglossus
megastoma, at Professor Herdman’s Fisheries Laboratory.

ADDITIONS TO L.M.B.C. COPEPODA. 103

Family LERN#IDA.

Anchorella appendiculata.
Several specimens were found attached to the gills of
the hake at Professor Herdman’s Fisheries Laboratory.

Fig.

Fig.

Sage eee te eee ee i

EXPLANATION OF PuaTE VI.
Pseudocyclopra stephordes, n.sp.

Adult male.
Anterior antenna.
Posterior antenna.
Mandible and palp.
Anterior foot-jaw.
Posterior foot-jaw.
Foot of first pair.

EXPLANATION OF PLATE VII.
Pseudocyclopia stephoides, n.sp.
Foot of second pair.
Foot of third pair.
Foot of fourth pair.
Fifth pair of feet, male.
Fifth pair of feet, female.
Abdomen and caudal stylets, female.
Abdomen and caudal stylets, male.

104,

ReEPoRT on the Investigations carried on in 1894
in connection with the LANCASHIRE SEA-FISHERIES
LABORATORY at University College, Liverpool.

By Professor W. A. Herpman, D.Sc., F.R.S., and
Mr. AnpREw Scott, Fisheries Assistant.

INTRODUCTORY.

Tus, the Third Annual Report of the Lancashire Sea-
Fisheries Laboratory, contains an account of the work
done during the past year, 1894, in endeavouring to
carry out the scheme of investigations drawn up by
Prof. Herdman when the Laboratory was established
in 1892, and is necessarily, to a considerable extent, a
continuation of what has already been published in the
two former Reports. One new line of enquiry this
year is the investigation, by means of ‘drift bottles,”
of the tidal and other currents in the Irish Sea which
would affect the distribution of small floating bod
such as fish eggs and surface food (see below).

Part of the work has been carried out on board the
fisheries steamer ‘‘John Fell,’ while making the usual
periodical visits to the in-shore and off-shore fishing
grounds of the district and adjacent parts of the Irish
Sea. The fishery officers, under the Committee, have
kept us supplied with material from their respective
districts. Owing to the resignation of two fishery
officers, this part of the work has been somewhat
interrupted during the latter part of the year, and,
consequently, the number of small fish sent in for
examination has been rather less than in former years.
This reduction in the number of fish is, however, counter-

SEA FISHERIES LABORATORY. 105

balanced by a great increase in the number of shell-
fish, z.e., cockles and mussels, sent in, as will be observed
from the statistics later on in this report.

The rest of the work, including the microscopical exam-
ination of the fish and shell-fish stomachs and eggs, has
been done in the Fisheries Laboratory at University
College, Liverpool. In March, 1894, Mr. Andrew Scott
succeeded Mr. Corbin as Fisheries Assistant, and the
detailed work during the year has been carried out by
Mr. Scott, under the general direction of Prof. Herdman,
as in former years.

Mr. Dawson has again kindly placed at our disposal
the monthly records of observations made in the district
by himself and the various fishery officers, and from
these we have extracted a good deal of valuable infor-
mation. Some of it is given here, while part is held
over for a future report in order that further additions
may be made before publication.

During the year Mr. Dawson has been continuing
the interesting experiments made with an ordinary
shrimp shank net fitted with a bar in order to determine
whether the number of small fish usually caught in shrimp-
ing could be diminished without affecting the quantity of
shrimps taken in the net, and, so far as the results go,
they seem to show that the bar shank net takes on the
whole fewer fish and more shrimps than either the
ordinary shank or shrimp trawl nets. We give the
results of some of the experiments further on.

A short account is given below of the Irish Sea as a
natural fishery ground, and the subject of a sea-fish
hatchery for the district is fully discussed. At the end of
the Report will be found some notes on a few additions to
the known fauna of the district obtained by the steamer
and otherwise during the trawling investigations.

106 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

We have to thank Mr. Robert A. Dawson, the Super-
intendent of the Fisheries, Captain Wignall, of the fishery
steamer, and the various fishery officers, for the willing
assistance rendered us in carrying out the various experi-
ments, and for supplying material for examination in the
laboratory.

EXAMINATION OF FooD IN STOMACHS.

In the course of the past twelve months 3,389 stomachs
of marine animals from different parts of the district
have been examined for the purpose of ascertaining what
forms the chief food of the animals in the area in which
we are more especially interested.

The following are the sources from which the stomachs
have been derived :— |

Food fishes up to three inches ............... 296

4 », above Ms 020.4 ee 1664
Other fish 8.9.42. Ce ee 67
Cockles, n2%).0 ht Ae Se ee ee — 629
Mie ele «techie tee HA et 0 ae pt ha 523
Slat eM S Ay ied ye ik ae ee ieee 290
Slaammlles 65 seb Asal Ae Ria Ve ee oe re 20

3389

THE Foop oF YouNnG FISHES.

From a further investigation into the food of the
young fishes in order to find out what forms the chief
source of nourishment after the contents of the yolk-sac
have been used up, it seems pretty certain now, as was
suggested in a former report, that after the larval fishes
have absorbed the food supply stored up in the yolk-sac, _
they pass to the stage in which copepoda form the chief
food, and when that stage is passed they take to feeding
on larger invertebrata, such as small annelids, mollusca
&C,

SEA FISHERIES LABORATORY. TOF

At the Scottish Fishery Board’s Marine Hatchery,*
Dunbar, some experiments were made last spring (1894)
by Mr. Harold Dannevig, in which he endeavoured to rear
young Plaice. He succeeded in keeping the newly hatched
Plaice alive in filtered sea-water till they attained the age
of 20 days by feeding them on material collected by tow-
net in Dunbar Harbour, but owing to stormy weather, the
water became loaded with fine mud, which was collected
in the tow-net along with the food-material and could not
be readily separated from it. This killed the young fish,
and so brought the experiment to an untimely end. An
examination of the tow-net material showed that it
consisted chiefly of Copepoda, in various stages of
development, and other young crustacea.

The following list gives the result of an examination
of the stomachs of 296 young fish from various parts of
the district. The list is divided up into five columns,
giving the locality, date, number of fishes examined,
range of size in inches, and the contents of the stomachs,
the limit of size being three inches in length. The size of
the smallest fish received during the year was half-an-inch,
they were collected at Lytham by Mr. Ascroft, who
kindly sent them to us, and an examination of the stomachs
showed them to be filled with the Copepod Hurytemora
affivis, Poppe. later on in the year we received a few
more small fish, also half-an-inch in length, from Capt.
Eccles, New Brighton. These had been caught near
Deposit Buoy, Horse Channel, Mersey entrance, but in
this case the stomachs were empty.

* An interesting description of the hatchery at Dunbar, and an
account of the very successful hatching operations carried on last spring
and summer, are given in Part III of the Twelfth Annual Report of the
Fishery Board for Scotland,

108 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Young Plaice (Plewronectes platessa).
| 13 Empty, 1 Cumacea,

4 Bi Gf
Morecambe. Dec. 41] 15 fish. | 12-8 inch. ib eis
9 Empty, 1 Copepoda,
Morecambe. Jan. 2, ee. 1 Annelida, 1 Coro-
phium.
‘ : 1 1 Empty,
Morecambe. Feb. 2| 3 5, |1¢-2 ,, 3 Animal Gees
9 Empty, 7 An. tiss.
y 7 Corophium,
Lytham. Mar. 16)| 28). 5 s|/2—=22 9 oe 3 Amphipoda (Bathy-
poreia) 2 Larval Crusta-
ceans, Ss
Blackpool. Mar. 21 | 5 4, |2<22 -,, 5 Empty.
Morecambe. Apts: £05) UW oy5 oe ei 1 Tellina.
Off Jumbo Buoy. pC Osa crea 2 ‘ 1 Mactra.
Deposit Buoy. Agr AS |i eye 25 a 1 Annelida.
Blackpool. Apr.<20))25 35, | 225.) es Empty, 3 Annelida,
17 Aninal tissue,
Lytham. May, 1S 45 aoe ne 8 Copepoda (Lurytemora 4
affiis). ™
Ay; 1 3 Annelida, 1 Amphi-
Bye LENA ge el ie seer poda, 3 Animal tissue.
Lytham May “9 5 3, 1125-2247, 2 Crangon, 3 Annelida.
Spencer’s Spit, May 11) 92> 5 22 5 2 Annelida
Mersey.
Deposit Buoy. May 235) 25, \3 33 2 Empty.
Lytham. May 30)" 4. 5925-0 95, 3 Empty, 1 Crangon.
Burbo Bank. Jume Qi 2 ete + 2 Empty.
Crosby Channel. | July 23} 2 ,, | 24 a 2 Empty.
Garston. Oct. alos! “Sie, 24 a 3 Empty.

4 Empty, 1 Annelida,

Barrow Channel. |Dec. 17} 6 ,, | 12-22 ,, 1 Crangon

Of the 132 young Plaice examined, 58 stomachs con-
tained no food, and 28 contained indistinguishable animal
matter, leaving 46 to be accounted for as having food. }

SEA FISHERIES LABORATORY. 109

Crustacea were found in 29 stomachs, or fully 63 %,
and consisted of Copepoda, Isopoda, Amphipoda, Cumacea,
and Crangon.

Annelida were found in 14 stomachs, or fully 30 %.

Mollusca were found in 2 stomachs, or fully 4 %.

Young Dabs (Pleuronectes limanda).
Morecambe | Feb. 2, I fish, 3 inch. | 1 Pandalus.

Lytham Mar 16)) i ,. | 24 ws 1 Empty.

4 Empty, 2 Copepoda
(Ectinosoma curticorne),
1 Copepoda (22 Eetino-
soma curticorne, 1 Longi-
Blackpool. Aor |) 105, | tLe, pedia), 1 Copepoda (Ee-
tinosoma, Thalestris, &
Canuella), 1 Cumacea
and Copepoda (Zetino-
soma), 1 Amphipoda.

2 Empty, 1 Copepoda

Blackpool. Apr. 20 Bo See ae as (Dera

Spencer’s Spit. May 11 ae ae: 7 1 Annelida.

Deposit Buoy. Ang. 21) 102 ,, |}4-24 .,, 86 Empty, 16 Annelida,
Ulverstone. Oct. 23 LT 555 | 2 e 7 aie (Jonestella
Garston. Dec. 14! 16 ,, | 24-22 ,, 16 Empty.

Of the 135 young Dabs examined, the stomachs of 109
were found to contain no food, the remaining 26 contained
food that could be recognised.

Annelida were found in 17 stomachs, or fully 65 %.

Crustacea were found in 9 stomachs, or fully 24 %, and
consisted of Copepoda.

Sprats (Clupea spratta).
1 Mysis, 1 Empty, 1

Lytham. | Mar. 16] 3 fish. | 2?—3 inch. Corophiun.

Blackpool. Mar. 21/6 ,, 2-22 ,, 6 Empty.

Morecainbe. Mar. 22]1 ,, 3 iP 1 Animal tissue.

110 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Sprats (Clupea spratta).

Deposit Buoy. Apr 182° 4. 3 as 2 Empty.

Leasowe Shore, | May 3/2 ,, 25 x 2 Empty.
Mersey.

Lytham. May 4/1 ,, 22 * 1 Empty.

Garston. Oct: 16/25 ,, 22-3 ,, 2 Empty.

Barrow Channel. | Dec. 17/1 ,, 24 - 1 Empty.

Garston. Dec, 14) 2 ,, | 2-3 3 2 Empty.

Soles (Solea vulgaris).
Morecambe. (Mar. 22/1 fish. [8 inch.

23»,

1 Cumacea,

Ulverstone. Oct. 2311 4, 1 Empty.

Raia maculata.

21 miles W.N.W.{ Apr. 3/1 fish. { 25 inch. -
from Morecambe

Bay Lt. Vessel.

1 Animal tissue.

Raia clavata.
21 miles W.N.W. | Apr. 3/ 1 fish.
from Morecambe

Bay Lt. Vessel.

22 inch. | 1 Empty.

Flounders (Pleuronectus flesus).

Grange-over- | May 28 | 3 fish. | 24 —22 inch. | 2 Empty,
Sands. 1 Crangon.

Yellow Gurnard (Trigla hirundo).
Deposit Buoy. | Sep. 18|2 fish, |2-24inch. | 2 Empty.

Foop oF LARGER FISHES.

Since the last Annual Report was published, we have
examined 1,664 food fishes, three and-a-half inches in
length and upwards, with a view to determine their food, _
and also the condition of the reproductive organs. These
fish were caught at various places both inside and beyond
the limits, as the table of localities will show, thus differ-
ing from the case of the young fishes examined, which

SEA FISHERIES LABORATORY. ae

were, with one or two exceptions, all from inside the
limits, and, therefore, as may be expected, it will be found
that there is a greater diversity of food in the stomachs.

Along with the food fishes, 67 other fish, not usually
looked upon as being suitable for food, have also been
examined, but their numbers are too small to be worthy
of record until they have been supplemented by the
collection of further material. These animals are of
considerable interest, however, from a point of view of fish
food, as they are frequently found in the stomachs of the
more valuable food fishes, so that it 1s of some importance
that we should find out what they themselves feed upon,
and to what extent they compete with the food fishes
feeding on the same ground.

As before, the statistics are arranged in columns showing
the locality, the date, the number of fish examined, the
size in inches, and the contents of the stomachs.

Plaice (Pleuronectes platessa).

Morecambe. Nov. 29] 2fish.|5£ inch. | 2 Empty.
Morecambe. Dec. 4/18 ,, | 34-6 ,, 17 Empty, 1 Crangon.
Morecambe. dam. Jt 7 5) ek —6 5, 7 Empty.
E x 1 Empty,
Morecambe. Jam, 24 )~2 5, [fatto 1 Cardium (Mollusca).
Morecambe. Hobs 2) 107 6,5 bod 9 5; 9 Empty, 1 An. tiss.
icc adi. 10 Empty, 1 An. tiss.
Morecambe. Feb. 8/12 ,, /83-6 ,, 1 remains of Crustacea,
3 miles below y 11 Empty, 2 An. tiss.
Lytham. Ber Ore PAG tee WWE ta 29 1 Jdotea and Cardium,
Morecambe. Feb. 20} 4 ., | 44-63 ,, 3 Empty, 1 An. tiss.
Bar ei 7 Annelida
Morecambe, Mar, 2} 9 5, |44-8 ,, oF ett BERS
OffMaughold Hd.,' Mar. 18 | 1g). \'104 2298 .. 15 Empty,

Isle of Man. |

3 Pecten tigrinus.

112 TRANSACTIONS LIVERPOOL BIOLOGICAL SOUIETY.

Plaice (Pleuronectes platessa).

‘The Hole. Mar. 14{ 2 fish. | 63
Lytham. Mar.16)15 ,, | 35—52
Blackpool. Mar.21|} 5 ,, | 62-84
Morecambe. Mar(229" 1°58 65
Morecambe. Apr. S1d4 4° | 4-105
North-West from | Apr. 1 ee re Pere

Jumbo Buoy.

Deposit Buoy ;

Horse Channel. Apr. 18] 6 4, | 33-7
Leasowe Shore. May 3/10 ,, | 384-92
Lytham. May 4| 2 ,, |7-83
Spencer’s Spit, May 11] 6 ,, | 34-63

Mersey.

Station IV, off

Blackpool. May NG) 5 53 1 Saul
Deposit Buoy, :

Horse Channel. May 23) 16 3; en = 8
Lytham. May 30/ 2 ,, | 384
1 mile east from|June 8| 6 ,, | 42-53

Newcome Knowl

Buoy, Mersey.

Burbo Channel. |June21| 7 ,, | 34-6
Grange-over- July 16} 3 ,, |4—52

Sands.

Crosby Channel. | July 23) 3 ,, | 384-74
Horse Channel. Aug. 9|29 ,, |9—138
Back of North | Aug.21/13 ,, |4

Bank, near De-

posit Buoy.

Deposit Buoy. Aug.28| 8 ,, |54—-7
4 miles West of ;
Blackpool. Aug. 31] 21 ,, | 9--163

inch.

39

2 Empty.

4 Corophium, 1 Crangon,
1 Crangon & Cardium,
1 Tellina, 8 An. tiss.

5 Empty.

1 Empty.

6 Empty, 6 TZellina, 1
Crangon, 1 sand only.

2 Mactra.

4 Empty,

2 Annelida.

9 Annelida, 1 Empty.

2 Cardium.

6 Annelida,

2 soft parts of Mollusca,
2 do. and Annelida.

3 Annelida.
3 Donax.

2 Empty.

6 Animal tissue. |

7 Empty.

3 Corophium.

3 Empty.

21 Scrobicularia, 2 Ani-
mal tissue, 6 Empty.

13 Mactra.

1 Annelida, 7 Mactra.

il Macrae
10 Mactra and Philine.

SEA FISHERIES LABORATORY. 113

Plaice (Plewronectes platessa).

3 Empty, 1 Nucula, 2
: Amphipoda(Ampelisca),
Off Bar Ship. Sept. 6/15 ,, |9-15 ,, 3 Mactra and Nucula, 4

Scrobicularia, 2 Mactra.

Off Blackpool. Sept. 6/10 ,, | 10—125,, 10 Annelida.

Deposit Buoy. Sept.18|27 ,, | 384-74 ,, pees aT, 3)
Burbo Bank. ce 4 ilk 35 t4—74 1 5, 11 Donax.

Garston. OGa16 | 7 4, lea 8t 5; 7 Empty.

Ulverstone Chan. | Oct. 23|53 ,, | 32-52 ,, 53 Empty.

Between Dingle| Dec. 14|33 ,, |4-9 ,, 33 Empty.

and Garston. .
Barrow Channel. | Dec. 17! 4 ,, |! 45-53 ,, 3 Empty, 1 Annelida.

- Of the 421 stomachs examined, 211 were empty and 23
contained indistinguishable animal matter, and one a little
sand only, leaving 186 which contained matter that could
be identified, and these are accounted for in the following
notes :—

Mollusca were found in 134 stomachs, or fully 72 %, and
comprised Cardiwm, Mactra, Donaz, Tellina, Scrobicu-
laria, Nucula, Philine, and Pecten. It will be observed
from the list that the majority of the fish with Mollusca
in their stomachs were taken in the Mersey district.

Annelida were found in 41 stomachs, or fully 22%; the
species were not identified owing to the mutilated condi-
tion of the specimens.

Crustacea were found in 15 stomachs, or fully 8 %, and
consisted of Crangon, Idotea, Corophium, and Ampelisca.

Thus, it will be seen that Mollusca occupy a very
prominent position in the food supply of the Plaice caught
in the Lancashire district, Annelida being second in im-
portance, and Crustacea third. This is almost the same
result as was arrived at in last year’s Report.

In the Firth of Forth, Annelida occupy the first place,

_— Mollusca the second, and Crustacea the fourth.

114 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Morecambe.

Morecambe,

Off Maughold Hd.,
Isle of Man.

The Hole.
Lytham.
14 miles W.N.W.

4 W. from More-
cambe BayLt.V.

21 miles W.N. W.
from Morecambe
Bay Lt. Vessel.

Morecambe.

Morecambe.

-Morecambe.

Deposit Buoy,
Horse Channel.

Leasowe Shore.

Spencer’s Spit.

12 miles W.N. W.
4 N. from More-
cambe Bay Lt. V.

14 miles N. by
W. from Liver-
-pool N.W. Lt.V.

Station IV, off
Blackpool.

10 miles W. 4 N.
from Morecambe
Bay Lt. Vessel.

Dab (Pleuronectes limanda).

Noy. 11
Mar. 2

Mar. 13

Mar. 14

Mar. 16

Apr. 3

Apr. 3

May 14

May 16

May 17

6 fish. 143-6 inch.

i

12

18

2?

99

93

id
.

6 ”
32-114,,
53-74 ”
5 29
45-10 ”
5— 62 > 7)
4—7 99”
34 =
43 »
34-53 9
Me) ae ae
45 — 84 Me)
64 -—104 ’9

He,

6 Empty.
1 Empty.

-|10 Empty,

1 Ophioglypha,
1 Pagurus and Pecten

6 Empty.

1 Empty.

8 Empty, 2 Annelida,
1 Doris, 2 Ophioglypha,
1 Ophioglypha & Monta-
cuta, 1 Solen, 1 Pagurus,
1 Animal tissue.

4 Empty.

1 Empty, 1 Amphipoda,
1 Empty.

1 Crangon.

5 Empty, 4 Annelida.

2 Crangon, 5 Annelida.

4 Annelida, 1 Portunus,
1 Solen.

14 Annelida,

2 Ophioglypha,
1 Pagurus,

1 Crangon.

5 Ophioglypha, 1 Anne-
lida, 2 Pagurus, 1 Mol-
lusca, 1 Zoophytes, 4
Empty.

2 Eimpty, 2 Ophioglypha,
1 Philine. ;

7 Empty, 18 Annelida, ~

|6 Ophioglypha, 2 Pa-

gurus, 1 Pagurus and
Solen, 4 Ophioglypha and
Solen, 4 Solen, 3 Ani-
mal tissue.

SEA FISHERIES LABORATORY. 115

Dab (Pleuronectes luomanda).

Deposit Buoy. May 23 ,

1 mile E. of New-

June 8
come Knowl] Buoy.

Burbo Channel. June 21

Crosby Channel. | July 23

Horse Channel. | Aug. 9

Back of N. Bank,
nr. Deposit Buoy.

Aug. 21

Deposit Buoy. Aug. 28

4 miles W. of

Blackpool. Slee

Off Bar Ship. Sept. 6

Deposit Buoy. Sept. 12

Burbo Bank. Oct. 4

Garston, Mersey. | Oct. 16

Between Dingle | Dec. 14
and Garston.

i)

43

24

15

15

16

29

299

d2

7)

9

1 Empty, 4 Crangon, 2
Annelida, 1 Annelida
and Crangon, 1 An. tiss.

1Empty,5Animal tissue.

3 Empty, 3 Crangon,
1 Annelida.

5 Empty, 3 Annelida.

4 Ophioglypha,

1 Scrobicularia& Mactra,
1 Mactra & Portunus,
1 Portunus, 1 Annelida,

6 Empty, 1 Mollusca
(soft parts),86 Annelida,

22 Empty, 1 Crangon,
1 Portunus.

6 Ophioglypha,
4 Philine, 5 Ophiogly-
pha and Philine.

| 2 Mactra, 1 Ophioglypha.

9 Empty, 4 Mactra, 1
Crangon, 1common shore
Crab (Carcinus).

4 Empty, 1 Crangon,
4 Donax and Nucula,

2 Empty.
16 Empty.

Of the 320 stomachs examined, 125 were empty, and
10 contained indistinguishable animal matter, leaving 185
to be accounted for as having matter in them that could
be identified, they are as follows :—

Annelida were found in 92 stomachs, or nearly 50 %.

Hehinoderms were found in 39 stomachs, or fully 21 %,
and consisted chiefly of Ophioglypha (sand stars).

Mollusca were found in 38 stomachs, or fully 20 %, and

_Ulverstone Chan. | Oct. 23| 1 , | 64. ¥9 1 Empty.

116 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

comprised Donaz, Nucula, Mactra, Philine, Scrobicularia,

Solen, Montacuta, and Doris.

Crustacea were found in 28 stomachs, or 15 %, and
consisted of Crangon, Portunus, Pagurus, and Amphi-
poda.

Zoophytes—One stomach contained some fragments.

The above notes show that Annelida appear to have
formed during the year a large per-centage of the food of
the Dab in our district. Hchinoderms, Mollusca, and
Crustacea also form a considerable item in the food supply.

Last year’s Report shows that the chief food during
the period dealt with in that Report, was Mollusca,
Crustacea and Annelida being next, but not nearly so
important as the first-named. :

In the Firth of Forth district, Crustacea form a large
part of the food supply, Echinoderms being second, Mol-

lusca third, and Annelida fourth. —

Flounder (Pleuronectes flesus).

Morecambe. Jan. 4{ 1 fish. | 6% re 1 Empty. .
7 Empty, 4 Annelida,
Morecambe. Jan. 24/12 ,, |4-82 ,,- |1 Crangon, and other
Crustacea.
The Hole. Mar.14] 2 ,, |94—92 ,, 2 Empty.
14milesN. by W.| Apr. 3} 2 ,, |10§-16,, 2 Empty.
from Morecambe
Bay Lt, Vessel. .
Grange-over- May 28| 2 ,, |4%—-5% 5, 1 Empty, 1 Crangon.
Sands.
Crosby Channel. | July 23 | 17 eS oy ae 10 Empty, 7 Crangon.
Horse Channely 9) Aus. 901 94, v.15 re 1 Annelida.
4 miles West of} Aug.31|} 1 ,, | 10 a 1 Mactra.

Blackpool.

SHA FISHERIES LABORATORY. 117

‘We do not give a per-centage statement of the food
of this fish owing to the small number examined, but
Annelida, Crustacea, and Mollusca appear to have been
the chief food of the flounder during the year.

From last year’s Report it will be seen that the order
was then somewhat different, Mollusca being first, Anne-
lida second and Crustacea and Zoophytes third.

Sole (Solea vulgaris).
Morecambe. Feb. 21; 2 fish. (5-5} inch. | 1 Empty, 1 Annelida.

3 Empty, 2 Cumacea,
Morecambe. Mar. 22| 9 ,, | 82-5 », | 2 Amphipoda, 1 Anne-
lida, | a little sand.

iamilesiy.N-W. | Apr. 3/10 .,, |e oes) | LOP Empty.
from Morecambe
Bay Lt. Vessel.

14 miles W.N.W.| Apr. 3) 2 ,, |134-164,, | 2 Empty.
4 W. from Liver-

pool N.W. Lt. V.

fommiles We by |Apr, 8| 9 5, | 9-15 » | 9 Empty.
N. from Liver-
pool N.W. Lt.V.
9 Empty, 1 Zoea of
Morecambe. Apr. 5\12 ,, |353—-55 ,, | Crab; 1 Cumacea, 1 other
Crustacean remains,

: 7 By as 12 Empty,
Morecambe. me WEIS 5, |ee—5 ” | 1 Ammphipoda,
Morecambe. Apr. 18] 5 ,, |42-6% ,, | 3 Empty, 2 Annelida.

Deposit Buoy, pr IS | 6 5, | 3s4—5% 4.” | 6 Empty.
Horse Channel.

Spencer’s Spit, May 5| 5 ,,.|4-42 ,, |1 Empty, 4 Annelida,
Mersey.

14 miles N. by | May 14/18 ,, | 83-19 ,, |17 Empty, 1 Solen.
W. from More-
cambe Bay Lt. V.

Deposit Buoy. May 23; 6 ,, | 4-8 », | 6 Animal tissue.
1lmile E. of New-|June 8| 5 ,, |44—54 ,, |5 Empty.

come Knowl

Buoy.

Burbo Channel. |June21|{ 9 ,, |44—74 ,, | 9 Empty.

— =

118 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

. Sole (Solea vulgaris).
Crosby Channel. | July 23 | 13 fish. |4—52 inch. | 11 Empty, 2 Annelida.

Back of N. Bank 4 Empty, 5 Annelida
nr. Deposit Buoy. ar) eel Crangon.

4 miles W. of | Aug.31| 4 ,, {114-138 ,, | 1 Empty, 3 Annelida,
Blackpool.

Off Blackpool. Sept. 6/12 ,, | 104-14 ,, | 12 Empty.
Heysham Lake. | Oct. 151 1 ,, [7 »» | 1 Crangon.

Of 151 stomachs of Soles examined, 115 contained no
food, 6 contained indistinguishable matter and one con-
tained a little sand only, leaving 29 which are accounted
for as follows :— :

Annelida were found in 18 stomachs, or fully 62 %.

Crustacea were found in 10 stomachs, or fully 34 % and
consisted of Crangon, Cuma, Amphipoda, etc.

Mollusca were found in one stomach only, or a little
over 38%, so that Annelida appear to be the chief source
of food for the sole, with Crustacea next in point of im-
portance.

Lemon Sole (Plewronectes microcephala).

14 miles W.N.W. | Apr. 3] 3 fish. | 74—82 inch. ] 2 Empty, 1 Annelida
3 W. from More-
cambe Bay Lt. V.

21 miles W. by| Apr. 3] 1 ,, |54 ,. |1 Empty.
N. from More-
cambe Bay Lt. V.

Morecambe. Apr. 18| 2 ,, |64-7 ,, |1 Empty, 7 Anneld?
Lytham May! 962° 46062-1115.) | 2Ammelidat

14 miles N. by| May 14] 2 ,, |8-11 », | 2 Annelida,

W. from Liver-

pool N.W. Lt.V.

10 miles W. 3 N.| May 17;17 ,, |54—-10 ,, | 6 Empty, 11 Annelida,

from Morecambe
Bay Lt. Vessel.

The food of this fish seems to consist chiefly of Annelida
but further data are required before we can state it
definitely. | ae

SHA FISHERIES

LABORATORY.

gs

In the Firth of Forth district, Annelda are also the
chief food, along with a few Crustacea and Mollusca.

Off Maughold Hd.,

Isle of Man.

Morecambe.

Morecambe.

Morecambe,

Morecambe.

Morecambe.

Morecambe.

The Hole.

Blackpool Closed
Ground.

Morecambe.

Back of N. Bank,
nr. Deposit Bnoy.

Garston, River

Mersey.

Brill (Rhombus levis).
25 inch.

Mar. 13

Apr. 25

1 fish.

1

bP)

43

1 fish (Whiting).

1 Crangon.

Turbot (Rhombus maximus).
| Nov. 29] 1 fish. |

4 inch.

| 1 Empty.

Cod (Gadus morrhua).

Nov.29| 4 fish. |4—53 inch.

Jane 4

Jan, 24

Mar. 14
Mar. 21

Apr. 18
Aug. 21

Oct. 16

11

9

oP)

44-6
43-74
4-6
193 - 27
93

5—6
43-54
7-8}

9?

1 Crangon. Sprat, and
Arenicola, 2 Crangon
and Gammarus, 1 Mysis
and Gammarus.

1 Empty, 2 Amphipoda
(Gammarus), 1 Crangon
and Annelida. 1 fish and
Mysis, 1 Portwnus.

1 Crangon and Mysis, 1
Crangon and Annelida,
1 Crangon and Shore
Crab, 1 Jdotea and My-
sis, 1 Gammarus, 1 Fish.

2 Empty, 1 Crangon,
1 Mysis, 1 Cumacean,
1 Annelida, 1 Mollusca
(Tellina), 1 Crangon,
Fish, and Amphipoda,
1 Crangon, Fish, and
Mysis, 1 Crangon, and
Mysis, 1 Mysis and
Cumacea.

2 Empty, 1 Fish.

1 Crangon.

2 Crangon.

2 Crangon

2 Crangon, 3 Pandalus,

120 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Cod (Gadus morrhua).
Ulverstone Chan. | Oct. 23| 2 fish. | 33 —44 inch. | 2 Crangon.

Between Dingle & | Dec. 14} 1 ,, |7 55 1 Empty.
Garston.

2 Mysis, 1 Mysis and

Barrow Channel. | Dec. 17 Amphipoda,

35, e| Seas” By,

Of the 46 stomachs examined 6 only were empty, the
remaining 40 being accounted for in the following notes :—

Crustacea were found in 36 stomachs, or 90 % and
consisted of Crangon, Pandalus, Mysis, Cuma, Amphipoda,
Idotea, Portunus and Carcinus.

Fish were found in 5 stomachs, or fully 12 %.

Annelida were found in 4 stomachs, or fully 10 %.

Mollusca were found in one only.

On comparing the above result with that given in last
year’s report it will be found to be almost the same.
_ In the Firth of Forth district, the food of the cod appears
to be very similar to that recorded for the Lancashire
Sea-Fisheries District.

Whiting (Gadus merlangus). |

. 1 Empty, 1 Crangon,
Morecambe. Nov. 29| 4 fish. | 34-3 inch. | 1 Crangon and Mysis, —
~ | 1 Amphipoda and Mysis.

2 Empty, 2 Mysis,

pecs, Dec. 4) 5 4, | 82-42 >> | 7 Cpustacean remains.
1 Amphipoda (Gamma-
Morecambe. Jame 45) 2 5 SSeS 4, Pe. doe. & Mysis.
Morecambe. anew eee 7e 5, | 2 Hmpty.
Morecambe. Feb. 2| 2 ,, |34-64 ,, | 1 Fish(Sprat),1 Empty.
2 Empty, 2 Cnn
Morecambe. Feb. 11} 9 ,, |4-6 4 2 Fish, | Cranganey

Amphipoda, 1 Annelida, |
1 Anime! tissue.

OffMaughold Hd.,| Mar.13| 4 ,, |7- 114 », | 4 Empty.
Isle of Man.

ay

The Hole, Mar. 14/26 ,, |74-20 ,, | 25 Empty, 1 Crangon.

Blackpool Closed
Ground.

Morecambe.

14 miles W.N.W.
s W. from More-
cambe Bay Lt. V.

21 miles W.N. W.
from Morecambe
* Bay Lt. Vessel.

16 miles W. & by

N. from Liver-
pool N.W. Lt.V.

Morecambe.

Morecambe.

Morecambe.

Lytham.

12 miles W.N.W.
from Morecambe
Bay Lt. Vessel.

145 milesW.N.W.
x W. from More-
cambe Bay Lt. V.

Spercer’s Spit,
Mersey.

14 miles W. 4 N.
from Morecainbe
Bay Lt. Vessel.

10 miles W. 4 N.
from Morecambe

Bay Lt. Vessel.
Deposit Buoy.

1 mile E. of New-

come Knowl Buoy.

SEA FISHERIES LABORATORY.

121

Whiting (Gadus merlangus).

Mar. 21

pI! 8

pre) 3

5 II

May 11

May 11

May 14

May 17

May 23

June 8

3 fish.

15

19

26

3

9)

4-5
38 - 62
aes:
74-11
Spel
”

31-5
44 —5
eo
7k = 98
BF = 10
Ad = 7
8h -11
Ok 11
44 - 6}
44 —~ 6

inch.

9)

9)

1 Empty, 1 Idotea,
1 Fish.

10 Empty, 3 Mysis, 1
Corophium, 1 Corophium
and Mysis, 1 Amphi-
poda, 1 Fish.

1 Empty, 1 Ophioglypha,

2 Crangon, 2 Fish (Sand
eels).

10 Empty, 2 Annelida,
2 Crangon, 1 Fish.

17 Empty, 1 Crangon
and Ophioglypha, 1 Por-
CUnus.

1 Corophiwm and Bathy-
pore.

4 Empty, 1 Fish, 1 Fish
and Crangon, 1 Mysis &
Gammarus, 1 Annelida.

1 Empty, 2 Crangon
1 Fish (Sprat).

2 Crangon

4 Empty.

14 Empty, 4 Pagurus,
3 Crangon, 1 Fish, 3
Amphipoda, i Annelida.

1 Animal tissue, 5 Fish.

3 Empty, 1 Ophioglypha,
1 Pagurus.

10 Empty, 2 Annelida,
2 Fish, 1 Crangon.

1 Empty, 3 Crangon,
2 Fish, 1 Amphipoda,

1 Empty, 2 Crangon.

saa aa ae a a id Tk Ea i EN No Ae lL a RRR a
ry “i, Fo a ee ’ - ‘ : ‘Coe

ree ok pili se pea at oe

OTs ae hae

~ Deposit Buoy. Sept.18 | 13

122 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Whiting (Gadus merlangus).

Burbo Channel. | June21]| 4 fish. {6-104 inch. , 2 Empty, 2 Crangon.
Crosby Channel. | June23|13 ,, | 44-8 », | 18 Crangon.
Back of N Bank. | Aug.21| 7 ,, | 84—5 » | 6 Empty, 1 Crangon.

» |4-64 5, | 13 Crangon.

Burbo Bank. Oct. 4/11 ,, | 44-54 ,, | 10 Empty, 1 Fish.
Heysham Lake. | Oct. 15} 5 ,, | 33-42 ,, | 8 Fish(Sprats),2 Empty.
Ulverstone Chan. | Oct. 283} 4 ,, | 4-5 ,, | 2Fish(Sprats),2 Empty.
Between Dingle| Dec. 14| 8 ,, | 33-8 > | & Empty.

and Garston.

Barrow Channel. ' Dec. 17° 2 ,, |4—8} 5, | 1 Isopoda,1 Amphipoda

Of the 261 stomachs examined 143 were empty, 2

contained matter that could not be distinguished, leaving

116 containing recognisable food; these are accounted for
in the following notes :— ;

Crustacea were found in 85 stomachs, or fully 73 % and
consisted of the following :—-Crangon, Mysis, Gammarus,
Pagurus, Idotea and Corophiwm.

Fish were found in 27 stomachs, or fully 24 %, these
consisted of sprats (Cluwpea spratta) sand-eels (Amvmodytes)..

Annelida were found in 7 stomachs, or nearly 7 %.

Echinoderms were found in 3 stomachs, being fully 2 %
and consisted chiefly of Ophioglypha.

In last year’s report the same order prevails with the
exception that the Echinoderms are replaced by Mollusca.

In the Firth of Forth district, the whiting appear to
feed largely on Fish, Crustacea being next in point of
importance, then Mollusca and Annelida.

Haddock (Gadus aeglefinus).
OffMaughold Hd.,, Mar.13, 4 fish. ,64—16 inch. ; 2 Empty, 2 Ophioglypha.
Isle of Man. :

The Hole. Mar. 14] 13 ,, | 64—114 ,, eke Sicha

SEA FISHERIES LABORATORY. 123

Haddock (Gadus aeglefinus).

14 miles W.N. W.
from Morecambe| Apr. 3] 4 ,, | 74-15 ,,
Bay Lt. Vessel.

2 Ophioglypha, 1 Por-
twnus, 1 Annelida,

21 miles W.N.W.
from Morecambe | Apr. 3| 1 ,, | 10 sy | & Crangon:
Bay Lt. Vessel.

144 miles W.N.W.
% N. from More-| May 11| 2 ,, |16—19 ,,
cambe BayLt.V.

1 Corystes, 1 Nucula and
Echinocyamus.

14 miles N. by W.
from liverpool| May 14} 1 ,, | 14 ne

1 Ophioglypha and Pec-
N.W. Lt. Vessel.

ten tigrinus.

10 miles W. 3 N.
from Morecambe} May 17] 6 ,, |8—14 3
Bay Lt. Vessel.

4 Empty, 1 Portunus,
1 Pagurus.

4 Mactra, 3 Scrobicu-
Off Bar Ship. pepe. 6) 9 -;- | 10-12) -,, | daria, “1. Annelida, 1
Annelida and Crangon.

3 Empty, 2 Annelids,
4 Mactra, 1 Annelida
Off Blackpool. Sept. 6/12 ,, |18-16 ,, | and Ophioglypha, 1 Mac-

tra and Scrobicularia,
1 Philine.

Of the 52 stomachs examined 19 were empty, the
remaining 33 being accounted for in the following notes :—

Mollusca were found in 18 stomachs, or fully 54 %, and
consisted of Mactra, Scrobicularia, Philine, Pecten, Nu-
cula and Solen.

Echinoderms were found in 7 stomachs, or fully 21 %,
and consisted chiefly of Ophioglypha, 1 stomach only
containing Hehinocyamus pusillus.

Annelida were found in 6 stomachs, or fully 18 %.

Crustacea were found in 6 stomachs, fully 18 %, and
consisted of Portunus, Corystes, Pagurus and Crangon.

Last year’s report gives the result of an examination of
37 stomachs from which it will be seen that Mollusca,
Crustacea, Annelida, and Hchinoderms were the chief food,
in the order given.

124 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

| In the Firth of Forth district, the Crustacea appear to
be the most important food of the haddock, then Mollusca,
5 Echinoderms and Annelida follow in the order named.

Herring (Clupea harengus).

Morecambe. Nov. 29| 1 fish. | 43 inch. | 1 Empty.

Deposit Buoy. Agar. 185) Awe. led eo ie 4 Empty.

Sprat (Clupea spratta).

= Morecambe. Nov. 25 | 2 fish. | 834-32? inch. | 2 Empty.
Morecambe. Jan. 4| 9 ,, |3%4-—43 ,, |.9 Empty.
Morecambe. Feb. 21} 2 ,, | 45-4 », | 2 Empty,
Lytham. Mar: 16) 5 4, sh— Sy | 5 Corophium.

Blackpool Closed Mare21 | 5 04. ik as as! 2 Empty,

Ground. | 8 Animal tissue.
Morecambe. Apr OU i. var Ppa fl! Empty.
a 6 Copepoda (Acartia,
ee Morecambe. | Apr. 5| 6 ,, |34—42 ,, Ge stage OE
mus.
Burbo Channel. | Jume Qi ay >, |44 39°) | Empty.

Between Dingle &! Dec. 14] 5 ,, | 385-33 ,, | 5 Empty.
Garston. *

Barrow Channel. |; Dec. 17} 2 ,, | 384-832 ,, | 2 Empty.

Grey Gurnard (Trigla gurnardus).

OffMaughold Hd.,| Mar. 13| 5 fish. | 7-138 inch. | 5 Empty.
Isle of Man.

a 14 miles W.N.W. =
| 2.W.from More-| Apr. 3| 6 ,, |6§-12 ,, : Bley a Fish,
cambe Bay Lt.V. : oe
; 21 miles W.N.W. 7 Empty, 25 Crangon,
from Morecambe | Apr. 3/39 ,, | 75-17 ,, | 6 Crangon & Fish (Sand
> Bay Lt. Vessel. eels), 1 Sand eels.
16 miles W. by
7 IN. “from: Tniver-WeApr, wo ae ae »,5 | Ll Crangon.
pool N. W. Lt. V

Deposit Buoy, Apr. 13| 2 ,, |3§-42 |», | 2 Crangon.

SEA FISHERIES LABORATORY. 125

Grey Gurnard (Trigla gurnardus).

2 Fish, 2 Fish and
» | 93-113 ,, | Crangon, 1 Crangon and
Mysis.

2 |
Station IV., off
Blackpool, Baye 10!) ®

12 miles W.N. W. AG etre: ‘
from Morecambe | May 11} 9 ,, |9-—104 ,, Fish. 2 iets ne

Bay Lt. Vessel.

14LmilesW.N.W.
EN, from More-|May 11| 5 ,, |94-114 ,, | 4 Empty, 1 Fish.
cambe Bay Lt. V.

14 miles N. by 2 Empty, 3 Crangon, 1
Mericom Wiver-|Mayl4|/ 7 ,, |8-10 », _| Amphipoda(A mpelisca),
pool N.W. Lt.V. 1 Ophioglypha.

31 Empty, 1 Annelida,
1 Mysis, 11 Fish, (Dra-
gonet and Sand eels), 1
Amphipoda, 4 Crangon.

12 miles W. 4 N.-
from Morecambe , May 17/49 ,, | 64-13 ,,
Bay Lt. Vessel.

Off Bar Ship. eps, or ie lo ») | 1 Crangon.

Off Blackpool. Depe Oe lyse pil 9,5 | 1 Crangon.

Of the 130 stomachs examined, 55 were empty, the
remaining 75 all contained recognisable food.

Crustacea were found in 55 stomachs, or fully 73 %,
and comprised Crangon, Mysis, Ampelisca, and other
Amphipods.

Fish were found in 26 stomachs, or fully 34 %.

Echinoderms were found in one stomach only, or a
little over 1 %.

Annelida were also found in one stomach only.

In the Firth of Forth district, the same conclusion holds
good as to the food of the Grey Gurnard.

Red Gurnard (T'rigla cuculus).

21 miles W.N.W. 1 Empty,
from Morecambe | Apr. 3] 3 fish. | 125-—134inch] 1 Crangon,
Bay Lt. Vessel. 1 Portunus.

1 Fish,

Off Lytham. May 9) 2 ,, | 18-163 ., | Hyas cearctatus.

ee Oe ae oe ee eee , - : Wee See io

126 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Red Gurnard (Trigla cuculus).

: 12 Empty, 1 Crangon &
| Portunus, 2 Portunus,

144 milesW.N.W. 4 Hyas coarctatus, 3

1N. from More- | May 1186 fish. | 8-15 inch. | ree a nd Hag 1
cambe Bay Lt. V. Amphipoda & Crangon,

3 Fish, 1 Galathea. 1
Annelida.

14 miles N. by W.
from Liverpool] May 14] 3 ,, | 74-10
N.W. Lt: Vessel.

», | 1 Empty, 2 Crangon.

9 Empty, 1 Portunus

10 miles W. 1 N. and Crangon, 7 Fish
from Monegan ne May 17; 21 ,, | 44-41% ,, Cpe! 2 Ss a ey
Bay Lt, Vessel. EOS, ta

1 Galathea & Portunus,
1 Crangon.
Off Bar Ship. Sep: 6! 2 ., 140 », | 2 Crangon.

Of the 67 stomachs examined, 23 were empty, the
remaining 44 all contained recognisable food.

Crustacea were found in 32 stomachs, or fully 72 %, and
consisted of Crangon, Portunus, Pagurus, Hyas coare-
tatus, Galathea, and Amphipoda.

Fish were found in 11 stomachs, or 25 %.

Annelida were found in 1 stomach only, or fully 2 %.

Yellow Gurnard (Trigla hirundo).
Spencer’s Spit. May 11| 6 fish. , 5 inch. | 6 Crangon.

14 miles W.4 N.| May 17) 4 ,, | 164-24 ,, | 4 Fish.
from Morecambe
Bay Lt, Vessel.

Deposit Buoy. May 23/ 5 ,, {43-52 ,, |1Empty, 4 Crangon.

1mile E.of New-|June 8| 3 ,, | 42-6 », | 2 Empty, 1 Fish.
come Knowl Buoy.

Burbo Channel. |June21] 2 ,, |7 », | 2 Empty.

Of the 20 stomachs examined, 5 were empty, the
remainder 15 contained recognisable food. ‘

Crustacea were found in 10 stomachs, or 50 %, and
consisted of Crangon only.

Fish were found in 5 stomachs, or 25 %.

SEA FISHERIES LABORATORY. 127

Grey Skate (Raia batis).

Blackpool Closed | Mar. 21| 1 fish. | 10 inch. | 1 Crangon.
Ground.

14 miles W.N.W. .
eWetrom Wore-| Apr. 3| 1 ,, | 34 = ee (Ampe:
cambe Bayt. V.

16 miles W. by
Heicom Iiver-| Apr. 3| 1 ,, | 14 », | 1 Empty.
pool N.W.Lt.V.

144 milesW.N.W. ZA Baapiy, 21) Bish.
4 N. from More-| May 11} 6 ,, |12-—26 ,, | Pagurus, 1 Portwnus, 1
cambe Bay Lt. V. Annelida.

14 miles W. by .

N. from Liver-| May 14} 1 ,, | 83 * eas phipoda (Ampe-
pool N. W. Lt.V.

Station IV, off| May 16| 2 ,, |14 15% ,, | 2 Empty.

Blackpool.

Deposit Buoy. May 23} 1 ,, | 4 ele Empty:

1 mile E.of New-|June 8} 2 ,, |7-74 ,, | 2 Crangon.

come Knowl Buoy.

Burbo Channel. |June21/ 1 ,, | 34 », | Ll Crangon.

Horse Channel. j Aug. 9} 1 ,, | 12 » | LyCrangon.

Peles West of| Aug.31| 5 ,, |95-17 ,, | 5 Crangon.

Blackpool.
Off Blackpool. Sept. 6/ 4 ,, |11-12 ,, | 4 Crangon.

Of the 26 stomachs examined, 6 were empty, the
remainder contained recognisable food.

Crustacea were found in 18 stomachs, or 90 %, and
consisted of Crangon, Portunus, Pagurus, and Ampelisca.

Fish were found in 1 stomach only, or 5 %.

Annelida were also found in | stomach, or 5 %.

Thornback Skate (Raza clavata).

Blackpool Closed ; Mar. 21| 2 fish. 8—94 inch. , 2 Crangon.
Ground.

OffManghold Hd.,| Mar. 31| 1 ,, | 27 3 |) empty.
Isle of Man.

14 miles W.N.W. ‘ dys
% W.trom More-| Apr. 3| 1 ,, | 11 FS ee dae). aoe Es

cambe Bay Lt. V. |

) 128 TRANSACTIONS LIVERPUOL BIOLOGICAL SOCIETY.

Thornback Skate (Rata clavata).

21 miles W.N.W. [2 Empty, ee
from Morecambe | Apr. 3] 5 ,, | 74-184 ,, a ee " 7 ee
Bay Lt. Vessel. (Sand Eel).

16muiles W. by N.| Apr. 3] 1 4, | 7 », | 1 Crangon.
from Liverpool

IN. W. lit. Vessel.

Lytham. May 9] 2 ,, | 144-17.,, | 1 Fish, 1 Pagurus.

Station IV, off 2 1 Crangon and Solen,
Blackpool. ee eee ee 5 Portunus and Solen.

14 miles W.N-W. | May 1|-3 ,,.9| 1720 «3% 2 Portunus, 1 Empty.
4.N. from More- a)!
cambe Bay Lt. V.

14 miles N. by W. - 2 Orangon, 1 Fish, Hyas
Hee Liverpool eae ee ee and Pagurus.

oW alt V.

14 miles W. 3 N. 1 Pagurus, 1 Nephrops,
from Morecambe | May 17| 4 ,, | 64-25 ,, |1 Portunus, 1 Crangon,
Bay Lt. Vessel. . {and Amphipoda.

Deposit Buoy. May 23:) 4 4, |3 », | 8 Crangon, 1 Empty.

1 mile EK. of New-| June 8! 4 ,, | 34-4 », | 4 Crangon.
come Knowl Buoy.

Burbo Channel. | June21} 8 ,, |4-42 », | 38 Crangon.
Back of N. Bank. | Aug. 21} 2 ,, | 54-6 »> | 2 Crangon.
Of. Blackpool. * “-Sept..6 | alos, “25 a 1 Lortins, e

and Crangon.

Of the 42 stomachs examined, 4 were empty, the
remaining 388 all contained recognisable food.

Crustacea were found in 36 stomachs, or fully 97 %, and
consisted of Crangon, Galathea, Portunus, Pagurus, Hyas,
Nephrops, Carcinus, and Amphipoda.

Moliusca were found in 6 stomachs, or nearly 16 %, and
consisted of Solen.:

Fish were found in 4 stomachs only, or 10 %. —

Sandy Ray (Rava maculata).

21 miles W.N.W.| Apr. 3] 1 fish. | 54 inch. | 1 Crangon.
, from Morecambe
, Bay Lt. Vessel.

Burbo Bank. Oct. 41 2 5, wi G4 =7 4, 2 Crangon.

SEA FISHERIES LABORATORY. 129

Cuckoo Ray (Rava circularis).

144 milesW.N.W.
4% N. from More- | May 11| 5 fish. | 6-14 inch.
cambe Bay Lt. V.

2 Empty, 1 Fish, 2 Am-
phipods (Ampelisca).

1 mile KE. of New. | June 8] 1 , 44 i 1 Crangon.
come Know] Buoy.

Back ot N. Bank.| Aug.21/ 1 ,, '6 “A 1 Crangon and Mysis.

The chief food of the various species of Skate seems to
be Crustacea and a few fish.
In the Firth of Forth district the same order prevails.

Sand Hel (Ammodytes lanceolatus).

Blackpool Closed | Mar. 21| 3 fish. ) 4¢—64 inch. | 3 Empty.
Ground.

Deposit Buoy. DIE Sr 2.) cis Si Gea gal” 12 Nishe
Burbo Channel. |June21| 1 ,, |.9 oe) el Banat

GENERAL CONCLUSIONS AS TO FooD oF FISHES.

On the whole, the results given in the above tables
bear out the conclusions arrived at in last year’s Report.
Although we shall go on with the examination of the
stomachs of fish as opportunity offers, it will probably
not be necessary to publish the detailed evidence in future
reports; we shall, therefore, in future give merely a
summary of the results, unless some unexpected and
novel facts turn up.

As to the practical application of the knowledge
acquired :—

Ist. It is of importance, as it enables us to ascertain
the characters of feeding grounds, so that we may
know whether a particular bay, bank, or tract
of off-shore ground is better fitted for one fish or
another.

2nd. It has a practical application in determining
whether the destruction of particular classes of

130 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. ‘

invertebrates, e.g., by a heavy trawl, has or has
not an injurious effect upon certain fishes.

3rd. When the time comes, as it probably will,
when it will be cheaper and surer to farm fish
than to hunt them, when fish are bred, reared,
and fed up for market, then fish food will have
to be accurately ascertained and carefully culti-
vated, and all such statistics as those we are now
accumulating will be of value and receive their
proper application. |

THE DISTRIBUTION OF IMMATURE AND OTHER FISH.

As already stated in the introduction, we have gone
carefully over Mr. Dawson’s statistics of the fish caught
in the district during the year, and we now give some of
the more important points which can be made out regard-
ing the distribution of immature and other fish.

Owing to the steamer having been largely engaged in
police work during the past year, there has not been so
much opportunity for trawling over the areas for which
we have given statistics in former reports, but what has
been done confirms, to a great extent, the figures already
given. ae

A continuation of the experimental trawlings on the
Blackpool closed ground seems to show that the number
of small fish that frequent this nursery 1s steadily increas-
ing, and thus fully justifies the closing of this area
against trawling, and we have no doubt whatever that if
some other parts of the district were dealt with similarly
the result would be equally satisfactory. Let us take,
for example, the Mersey. From the middle of October
till about the end of December, there are great numbers
of young fish’in the river between the Dingle and
Garston, chiefly Plaice and Dab, 2+ inches and upwards,

SEA FISHERIES LABORATORY. 13t

very few of them exceeding 9 or 11 inches in length, and
as the ground is frequently trawled over both with shrimp
and ordinary trawls, the destruction of small fish is very
ereat. After the middle of December the fish begin to
leave this part of the Mersey and come down to the banks
off Crosby and Formby. These banks are then in their
turn continually trawled over, so that the destruction goes
on for a large part of the year. If it 1s not possible to
apply to this part of the district the same thorough
measure which has been applied to Blackpool, and entirely
prohibit trawling in any form, a close time, which would
take effect when the greatest number of fish are present
on the respective areas, would be very useful, such as
that the upper parts of the Mersey be closed against
trawling from the beginning of October till the end of
December, and that the lower parts be closed from
January till some time in summer. ‘There seems to
be especially heavy destruction in July, August, and
September around Burbo Bank and near the deposit
eround. ‘The experiment might be tried for a couple of
years to ascertain what amount of beneficial effect would
result from a partial closure, statistics being collected
weekly or monthly. So long as no artificial hatching is
done in the district, it is only by a most careful protection
of the natural rearig grounds of the young fish that the
quality and quantity of the larger fish on the various
fishing grounds around our coasts can be maintained.
As to the necessity of supplementing these restrictive
measures by hatching, and so adding to the supplies in
the sea, that is discussed elsewhere (see p. 136).

In the following list we give the average number of
fish caught on the Blackpool closed ground during the
year, and for the sake of comparison we also give the
figures for the previous year from the last Report :—

132 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

1892. December 2471 1893. December 20889*

1893. January 134 1894. January 8512*
February 1199 February 4729*
March 638 March 3896
April 302 April 1688*
May 101 May 802
June 122 June 1440
July 385 July 1204
August 437 August 3362*
September 14892 September ==
October — October —
November 2536 November 10810

The figures marked thus * are single hauls only, and in
the months left blank the pressure of police work pre-
vented any experimental hauls being taken. ‘Taking the
results of 1894 as a whole it will be seen that there is a
marked increase on the number caught during the former
year, and they also bear out the remarks made in last
Report as to the months in which the greatest and
smallest number of fish are present on this ground. It
appears from the figures now before us that the number
of fish on this particular part of the district reaches its
maximum in December, and then eradually goes down

_ till the month of May, after that it begins to increase, and

goes on increasing till it again reaches the maximum.

As in last year’s Report we again give a comparison
between the numbers of fish taken on the Blackpool ground
and those taken in Morecambe Bay, the estuary of the
Mersey, and in the Ribble estuary, the areas included in
these districts being the same as those given in the Report
for 1893 at page 86.

Each column contains the average number of fish
caught with the shrimp trawl, the figures marked * being

single hauls only.

Mersey. Ribble. Blackpool. Morecambe.
1893. December 593 — 20889* 231
1894. January 507 1950 8512* 131

SHA FISHERIES LABORATORY. so

1894, February 492 _ 4729* 197
March 871 -— 3896 220*
April 378 — 1688* 91
May 828 = 302 186
June 686 ~~ 1440 57
July 1186 as 1204 1533
August 3149 — 3362* 2250
September 2118 — — 2200
October 1331 — — 2350
November — — 10810 2150

These columns also bear out the remarks made in last
year’s Report as to the time when the fish seem to reach
the maximum and minimum limits.

VITALITY EXPERIMENTS.

The following are the vitality experiments which have
been made during the year by Mr. Dawson on board the
steamer :—

I. December 28th, 1893. Put in tank from Black-
- pool closed ground—
38 Soles, 3 inches in length,
15 Plaice, from 13 to 7 inches in length,
4 Dabs, from 13 to 3 inches in length,
After being in tank 34 hours—

1 Plaice, 14 inches long, and 1 Dab, 2 inches
long, were dead, the remainder were very
lively.

II. March 8th, 1894. Blackpool closed ground—

2 Soles, 3 inches in length,

6 Plaice, 3 to 74 inches in length,

4 Dabs, 33 to 74 inches in length,

After being in tank 2 hours and 5 minutes, all the
fish were alive.
III. March 19th, 1894. Blackpool closed ground—

12 Plaice, 2 to 6 inches in length,

12 Dabs, 4 to 6 inches in length,

134 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. ‘

4 Codling, 4 to 5 inches in length,

All the fish appeared lively with the exception of _ “

2 Plaice, 2 inches in length, and 2 Codling, 4
inches in length, which appeared deadly. |
After being in tank 5 hours and 10 minutes, all the
fish were alive (very lively).

IV. March 23rd, 1894. Shrimping ground, near 4

Deposit Buoy, Mersey—
13 Soles, 33 to 7 inches in length,
6 Plaice, 23 to 6 inches in length,
2 Whiting, 6 inches in length,

The fish had been lying on the deck and in the
sun for 8 minutes before being put in the tank.

Flat fish appeared lively and the round fish

deadly. One of the Soles was left amongst the
other fish and debris in the basket for 20 minutes

before being put in the tank among the others.
After being in tank 2 hours, one of the Whiting —

was dead owing to an injury received to the air-
bladder; the remainder were lively, and were ~

returned to the sea.

V. March 23rd, 1894. Shrimping ground, near =

Sag

Deposit Buoy, Mersey—
12 Plaice, 23 to 5 inches in length,
12 Dabs, 4 to 5 inches in length,
6 Soles, 3+ to 4% inches in length,
All the fish had been lying in a basket in the sun
for 20 minutes before being put in the tank.

After being in the tank for 13 hours all the fish

were lively. :
VI. April 2nd, 1894. Blackpool closed ground—
bia 12 Plaice, 24 to 7 inches in length,
12 Soles, 3 to 53 inches in length,
These fish had been left lying in a heap ina basket

SEA FISHERIES LABORATORY. $35

amongst other fish for } an hour before being
put in the tank and for the most part appeared
deadly.

At the end of three hours all the fish with the
exception of three Soles were alive, but two of
these had been killed by being jammed in the
discharge pipe.

VII. April 6th, 1894. Deposit ground, Mersey—
12 Plaice, 5 inches long,
12 Dabs, 5 inches long,
6 Soles, 44 inches long,
6 Whiting, 6 inches long.

A number of the fish appeared deadly when put in
the tank. After being in the tank 5 hours all
the fish with the exception of 1 Whiting were
very lively.

VIII. April 30th, 1894. Off Jumbo Buoy, Southport
entrance—
12 Plaice, 4 to 6 inches long,
8 Dabs, 4 to 6 inches long,
1 Sole, 7 inches long,

After being in the tank 2 hours all the fish were
lively.

IX. May Ist, 1894. Near Jumbo Buoy—
12 Plaice, 6 inches long,
12 Dabs, 7 inches long,

After being in the tank 2} hours all the fish with
the exception of one Dab were lively.

X. May llth, 1894. 12} miles W.N.W. from

Morecambe Bay light vessel—

At 7 p.m. (Friday) put 4 Soles 9 to 15 inches long
into the tank and another Sole from the next
haul. At 12 midnight transferred the Soles from
tank into a tub having a number of holes bored

1386 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

in it, then put tub overboard in Douglas Harbour.

At 9.80 a.m. (Saturday morning) all the fish were © |

alive and very lively, but at 8.30 in the evening
2 were dead and at 8 a.m. on Sunday morning
only one remained alive, the others having died.
During the time the tub had been in the water
it had been constantly oscillating.

XI. May 14th, 1894. 10 miles W.2N. from More-
cambe Bay light vessel—
At 9.45 a.m. put 6 Soles, 10 to 16 inches, into the

tank. 9 more were added at 12.45 pm. and
Spams |

At 6 p.m. the circulation had to be stopped owing _

to the steamer going into fresh water, the soles
being then all alive and very lively.
XII. May 16th, 1894. Blackpool closed ground—
12 Soles, 33 to 5 inches long, |
5 Plaice, 24 to 5 inches long,

6 Dab 33 to 5 inches long,

After being in tank 84 hours all the fish were alive.
These experiments lead to the same conclusion as those
we reported upon last year, viz., that under ordinary —
circumstances the great majority of the fish taken would
survive if returned at once to the sea. Ground-frequenting
fish of sedentary habits probably stand exposure best, and
soles seem to be especially hardy in this respect. The
practical application of these experiments is that 1t is well
worth while taking some trouble to ensure that all under-
sized fish brought up in the trawl should be returned to
the sea as quickly as possible.

Fish CULTURE AND HATCHING.

This northern area of the Irish Sea, in the centre of
which the Isle of Man lies, and which contains our Lan-

SEA FISHERIES LABORATORY. 137

cashire Sea-Fisheries district, is itself one large natural
sea-fish area, with its own spawning grounds, nurseries,
and feeding grounds, independent, so far as the greater
part of its fish population goes, of neighbouring seas,
but having its inshore and offshore grounds interdepen-
dent and intimately connected with one another, by the
successive stages and migrations in the life histories of
the Food fishes. Consequently it is most unfortunate
that our national and international laws are such that the
area cannot be treated as a compact whole each part of
which is of importance in the interests of the fishing
industries. The whole of the Irish Sea ought to be under
the jurisdiction of one authority, so that fish may be
protected, when necessary, in any part of it, so that the
same bye-laws may, if required, apply to Lancashire,
Anglesey, and the Isle of Man, and so that, to take a
particular case, the Sole may be protected when spawning
in the deep water of the offshore grounds. The three mile
limit whatever it may be from the point of view of national
defence and international arrangement, 1s an absurdity
from the fishery point of view, and all efforts to improve
the fisheries jn a district like ours are severely handicapped
by the fact that the fish, their enemies, their food, and
their captors can so readily pass beyond the range of all
regulations. We can doa good deal, it is true, by prevent-
ing the destruction of young fishes in the shallow waters
round the coast, but we cannot do nearly as much as is
desirable so long as no protection can be afforded to the
fish when spawning on the offshore grounds.

Excepting such a case as the Herring, a migrating fish
with demersal eges, the life history of one of our typical
food fishes is probably much as follows:—During the
ereater part of the year the adult fish moves from place
to place throughout the district, being influenced in its

=—— “+t | oe

188 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

wanderings chiefly by the search for food. As a general
rule the Sole, for example, is feeding in winter in the deeper
waters of the offshore grounds, while in spring the smaller
ones at least come in again to the shallower water off the
banks. At the spawning season, in the case of the Sole
from May to July, the large fish go away from land and
congregate on the spawning grounds which in this district
(as was pointed out in last year’s report) are tracts of —
deeper water, 20 to 25 fathoms, lying off the east of the
Isle of Man, or in the central part of the Irish Sea. There
the ova are extruded and fertilized, and then rise as young
embryos to near the surface of the sea. They are now
at the mercy of the winds and waves, they are carried in
various directions (see section on surface currents, p. 154),
and are exposed to the attacks of innumerable enemies,
including most of the animals which live around them in
the sea and even their fellow fish. Those that survive are
eradually carried by tidal currents into the shallower |
waters round the coast, and here the young fish, which
have now passed through their embryonic and larval
stages, leave off their pelagic mode of existence and take

to bottom feeding. It is at this stage that they make

their appearance (in April, May, and onwards in the
summer) in the fish nurseries round our shallow sandy
coasts, and there they live and grow, with occasional
wanderings determined by season and food, until they
reach the adult condition. In the warmest part of summer
they are frequently found moving up the estuaries (in
August, September and October this year Plaice and
Haddock have apparently been present in great abundance
off the Ribble estuary), and then with the first cold weather _
they move out of the estuaries to the banks.
Consequently it is evident that the supply of the inshore
nurseries depends upon the preservation of the offshore

SEA FISHERIES LABORATORY. 139

spawning grounds. If then, it is impossible at present to
take any steps for the protection of the old fish when
spawning, can anything be done to ensure a more abundant
supply of larval fishes to stock our bays? The only help,
in addition to protection, that can be given by man to the
fish-population of an area is by artificial fish culture and
hatching. On account of the enormous numbers of ova
which food fish produce it is possible for man, by stepping
in and saving even a minute fractional per-centage from
the destruction which normally takes place, to mcrease
very largely the number of young in a given area of the
coast waters.

These considerations shew the importance of sea-fish
hatching, and in an area like the Irish Sea there ought to
be, and one can scarcely doubt but that there will soon
be, a central fish hatching and rearing establishment in
some convenient spot where the water is the purest
obtainable and as free as possible from all suspended
matters. Such a hatchery should not be for the benefit
of Lancashire alone, nor of Cheshire, nor of the Isle of
Man. The locality ought to be chosen on the merits of
its physical conditions alone, apart from any ideas of
County boundaries and limits of jurisdiction of local
authorities. In a previous report, Port Erin at the 8.W.
corner of the Isle of Man was recommended for this
purpose, solely because of the known purity and clearness
of the sea water on that rocky coast, because of the
proximity to the spawning grounds of the most esteemed
fish, and because of the presence of the Biological Station
at Port Erin from which assistance and advice might
readily be obtained in regard to many of the difficulties
which would probably turn up during the first few years
of working. Spawning fish could be obtained in the
season either from the trawlers at Port Erin or from the

140 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

‘“‘ John Fell,” and the fry when hatched could be conveyed
to any selected spots on the coasts of Lancashire, Cheshire,
or North Wales and set free on suitable ground.

Port Erin is probably not the only suitable spot for a
hatchery in the district, but it 1s the one where the physical
conditions are best known. ‘There may be other places
on the coast of the Isle of Man, or in North Wales,
possibly even in Cumberland, which are equally suitable
for the establishment of a hatchery, but they have still to
be investigated. Port Erin and its water, the temperatures
and specific gravities, the fauna and flora, and all the
surrounding conditions are pretty well known to us. In
regard to the other coasts of our district, Cheshire and
Lancashire, the shallow waters around them although
apparently admirably suited as nurseries for fish in some-
what older stages, are so exceedingly muddy that it would
be a very difficult and probably expensive matter to render
them sufficiently free from all suspended particles as to
be fit for hatching purposes. The elaborate precautions
that have to be taken even at Dunbar in order that the
water may be sufficiently filtered bring forcibly before one
the inestimable advantage of having a pure supply of
water to begin with.

We have paid some attention lately to the water around
Piel Island, in the Barrow Channel, as that spot has been
proposed as the site of a hatchery for Lancashire. On
two special visits, along with Mr. Dawson and others, in ~
the Fisheries Steamer, the specific gravity, temperature,
and living contents of the water were examined, and since
then some samples of water taken by the bailiff at Piel
have been sent to the Laboratory for examination. On ~-
June 5th, 1894, the specific gravity in different parts of
the Channel from Barrow to Piel Island varied from 1:026
to 1.027, the temperature of the bottom water was 54° F.,

SEA FISHERIES LABORATORY. 141

and the tow-nets showed plenty of ordinary surface forms,
both diatoms and animals, medusoids being especially
abundant. All this is satisfactory enough, and the water
sent to the laboratory by the bailiff has been of good
quality, although the sp. gr. has been only about 1°022.
In looking at the sea off Piel Island however the water is
seen to contain much fine suspended matter, this is
especially the case when one examines it close to shore
from a small boat, and when wading along the edge at
low tide. No doubt it would be possible to filter such
water, but the process would, judging from what has been
done at Dunbar, be troublesome and expensive and any
temporary breakdown in the filtering arrangements would
be fatal. It is giving the experiment—for the first year of
working of a sea-fish hatchery in a new district must be
more or less experimental—a poorer chance of success to
handicap it with water which is not the cleanest obtainable
in the Irish Sea.

Although at the time when the specific gravity was
taken, off Piel Island, on June 5th, the salinity of the
water was perfectly satisfactory, still 11 was much lower
when tested in November and December and there is
some reason to doubt—judging from the great variations
in the specific gravity observed in places off the Lancashire
coast (e.g.,1n Crosby Channel our statistics range from
1018 to 1:026)—whether it would remain sufficiently
constant for hatching purposes. In one of his earlier
reports on sea-fish hatching in Norway, Captain Dannevig
complains of the large proportion of deaths due to varia-
tions in the specific gravity, and furthermore we notice
that during the remarkably successful first season’s work
at Dunbar last year, when about 27 millions of young
plaice and half-a-million of young cod were hatched out
and set free on the Scottish fishing grounds, with a loss

ee ee ae
a :

142 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of only 4-4 per cent, the water supplied to the hatchery
was almost invariably at the normal specific gravity 1:0270,
only once sinking to 1:0264. ,

In the event of the salinity at Piel remaining fairly
constant, and provided that the water can be successfully
filtered, a hatchery placed there would in all probability be
a success, and the same might be said of Hoylake which
has also been suggested as a site for a hatchery, but in
both cases the initial difficulty, the muddiness of the water,
has still to be overcome. Hence we are led to the con-
clusion that in place of a hatchery for Lancashire being
established at Piel Island and one for Cheshire at Hoylake
it would be much better if the various County Councils
and Fishery Boards* having jurisdiction over the terri-
torial waters of the Irish sea could see their way to unite
in promoting a fish hatchery at Port Erin for the benefit
of the entire area.

In concluding this section of the report we desire to
re-state what we regard as one of the most fundamental
points which can come before the consideration of a Sea-
Fisheries Committee: viz., that there are two methods by
which the decadence of a fishery may be checked, (1) by —
killing fewer undersized fish, and (2) by adding to the
total number of fish living in the district. Consequently
it is most desirable and important that Sea-Fishery
Committees all round the coast should, in addition to any
restrictive legislation that may be required, undertake
directly productive action such as sea-fish hatching and
rearing, and aquiculture in general.

* A measure has passed the House of Keys and has lately been promulgated
conferring powers to make fishery bye-laws and other regulations upon a
Committee in the Isle of Man.

SHA FISHERIES LABORATORY. | 143

SHELL-FISH.
I. MUSSELS.

During the year 523 mussels (Wytilus edulis) have been
examined with the view of finding out the time of spawn-
ing and also the food of this important shell-fish. So far
the result of this examination seems to show that the
food of the mussel consists of Diatoms, Spores of Aleve,
Vegetable debris, Foraminifera, and remains of Copepoda,
all of which are no doubt brought within reach of the
mussel’s ciliary currents by the motion of the surrounding
sea water; the variety in the food will therefore depend to
a very large extent upon what is the general fauna of the
mussel bed. As the food is, however, comparatively well
known now, our attention was more especially directed
to the determination of the spawning period, and as far as
the year’s results go they appear to show that the mussel
reaches maturity about the middle of May, and that the
spawning period lasts to the middle of July. There seem
to be a few exceptional cases, but the majority of the
mussels on the Lancashire coast have probably finished
spawning by the time stated.

In a report in 1886 by Mr. John Wilson, B.Sc., on the
development of the common mussel, published in the
appendix to the Fourth Annual Report of the Fishery
Board for Scotland, p. 218, it is stated that ‘‘ Professor
M‘Intosh found that in general the mussels reached full
reproductive maturity in April; thereafter the ova and
spermatozoa gradually disappeared from the mantle, until
in July those he examined were spent.”

The following list gives the results of the examination

of the samples sent to us.

. 4Empty,7 Spores,
3 female | nearly ripe. | Diatoms and Veg-

11 fish. ale
Smale f etable debris.

Morecambe. Jan. 24 2—24 inch.

144 TRANSACTIONS LIVERPOOL

Fleetwood.

St. Anne’s, lower
Scar.

St.Anne’s, Light-
house Scar.

Lytham, Church
Scar.

Morecambe.

Morecambe.

Grange over Sands
Humphrey Head.

Lytham; 1 mile
belowSt. Anne’s.

Morecambe.

Morecambe,
Heysham Scars.

LythamSt.Anne’s.

Lytham,nearPier.

Roe Island.

Lytham.2 yrs. old
from St. Anne’ Ss
beds.

Lytham, Church
Scar.

Lytham, Formby
Sear.

Roe Island.

Jana

Feb.

pot

Feb.

jp

Feb.

_

Reb. 2

Mar. 22

Apr. 10

Apr. 10

Apr. i

Apr. 11

Apr. 18

Apr. 18 |

May 1

May 3

May 10

May 22

{| May 24

LN)

‘e
~~

BIOLOGICAL SOCIETY.

9 female
4 male

1 female |
lmale }j

4 female
8 male

8 female

eas jalmost ripe.

3 female) __ en
a }near y vipe.

6 female

5 ae } immature.

12 male, immature.

4 female \ about 4
3 male mature.

6 female A

6 whale immature.
7 female \ . ea
4 male immature.
5 female) about 4

1 male mature.

5 feral, ova ‘07 m.

5 male sperms moving.

5 female ova '06-07m.

7 male, sperms not
moving.

10 female, ova ‘07 m.
2 male sperms moving.

2 female, ova 07 m.
6 male, sperms not
moving.

3 female, ova ‘09 m.

4 male, sperms not
moving.

7 female, ova °09

5 male sperms moving.

} almost ripe.
almost ripe.

}almost ripe.

4 Empty, 9 Spore . ? .
Diatoms and Veg-
etable debris.

2 Spores, Diatoms _ .
and Vegetable den
bris. 4
12 Spores, Dae
toms and Vegeta- a
ble debris. x

16 Spores, Dia-
toms and Vegeta-
ble debris. ;

8 Empty.

14 A little mud.
6Empty,6Spores, — =
Diatoms and Veg-
etable debris. ~

7 Spores. a

7 Empty, 5 sand —
and Vegetable
debris. tA

11 Empty.

6 Spores and Veg-
etable debris.
10 Sporn and V eg-
etable debris.

7 Empty, 5 Spar
Diatoms and Veg- —
etable debris.

2 Empty,10Spores 4
and Vegetable
debris.

8 Mud and
Diatoms.

3 Empty,
4 a little mud.

11 Empty, 1 sand
and Vegetable
debris.

|
Grange overSands| May 21

Grange over Sands

Humphrey Head. ae 2°
Lytham. May 30
Lytham,

Horse Bank. June 6
Grange over Sands

Waded Scar. J
Duddon Channel. | June 21
Roe Island,

Rusebeck Scars. Tea
Lytham, St. Anne's
- Mussel beds. ey 2
Barrow Channel. | July 18
eee Pe Anne's July 19
Grange over Sands

Humphrey Head. July 25
a> Anne's July 24
Duddon Channel | July 31
Roe Island, Ruse-

beck outer Scar. eae 6
Grange over Sands Sy

UlverstoneBridge ane!
ames Aug. 7
Grange over Sands

Humphrey Head. oro
Roe Island, oie

Rusebeck Scar. | AY: 78

SHA

12

12

12

14

12

16

10

10

10

12

12

12

FISHERIES LABORATORY.

29

fish.

Pais
2-93
Qh 94
igs
12-2
2-24
2

23-3
2

De = Bt
12 - 24
2-28
2-24
2-24
ie
24 - 24
2-24
2- 24

?

4 female, ova ‘1 m.

8 male, sperms not
moving.

9 female, ova ‘07-*1m.

3 male, sperms not
moving.

5 female, ova ‘09 m.

7 male, sperms not
moving.

5 female, ova ‘09 m.

9 male, sperms not
moving.

4 female, ova ‘09 m.

8 male, sperms not
moving.

4 female, ova ‘1 m.

4 male, sperms not
moving.

12 male, sperms not
moving.

12 ? spent.
12malesperms moving

16 ? spent.

7 ? spent.

10 ? spent.

3 female, ova ‘07 m.
7 male, sperms not
moving.

10 ? spent.

4 female, ova *12 m.
4male ?

12 spent.

12 spent.

12 spent.

145

12 Empty.

7 Empty, 5 Spores
and Vegetable
debris.

12 Empty.

14 Empty.

12 Empty.

8 Empty.

12 Empty.
12 Empty.
12 Empty.
16 Empty.
7 Empty.

10 Empty.
10 Empty.
10 Empty.
8 Empty.

12 Empty.

1 Empty, 11 Vege-
tabledebris&sand

12 Sand, Diatoms
and mud.

TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. F

146

Duddon Sands. Sept.18|/10 ,, | 24-24 ,, | 10 spent. 10 Empty. %

Roe Island, , 1 A little
Bs : - Waded Scar. Sept.19| 7 ,, | 23 peel Sem: 6 Empty. aioe
— Baicliff, ales 1 Empty,

West Hollow. Pepi 2O ee 2 ee age Cea 8 Sand and spores

Grange over Sands Qo iso mal 9 Empty, 2 Vege: :

Spe Cee ae 11 spent.

|

|
a - HumphreyHead. | table debris.
? A Ulverstone Chan- 7 Empt , 5 Vege-
e* nel, WestHollow. Oct. 11/12 4, | 23722 ., | 12 spent. table eae e |
ot: ih
a Barrow Channel. | Oct. 15) 7 ,, | 24-24 ,, | 7 immature. 7 Empty.

3 female) very
: , 195 . .
2 Grange overSands| Oct. 16;12 , | 24-28 ,, ny i immature, | {22 eae

Grange overSands | 4, Tey 1 female, ova ‘12 m. | 10 Empty, 2Vege-
Humphrey Head. eet ae eps Talae pee ep 11 immature. table debris. be
7 3 female, ova ‘1m. | 4 Empty, 6 Vege a

me Roe Island
Gor 0) 10 7 immature. . table debris.

Rusebuck Sear.

Grange overSands| 7g | 19 9.86)

4 female, ova ‘1m. | 10 Empty, 2Ve; ge-
Waded Scar. e

> | 8 immature. table debris.
Morecambe Chan- Nov. 12 | 19 e.

9 female, ova ‘1 m. 11 Empty 1Veg
nel, deep water. | 19-23 ,

ao 3 immature. table debris. —

ig 2 female alle :
“— Eee i
a oo Docks. Nov201 58 e,) zee : ne immature. | 8 Empty. P

eee
7

Humphrey Head. | Dec. 4 | 138 ,, |15-23 ,, : ela’ } immature. Vegetal fees a

Our mussel fisheries, valuable as they are, might, we a
have no doubt, be made still more valuable by the intro-
, duction of some artificial cultivation. Our coast waters are
. well supphed at present with mussel seed (the free-swiming
embryos), they are found year after year settling down on
all sorts of suitable and unsuitable submerged objects, and
many millions of them every year perish miserably for
want of a little looking after. As was recommended in
last year’s report, simple stakes (bouchots without wattling)
should be erected in likely spots, not for the purpose of
taking the place of the beds—which are probably better _

“a
suited on the whole to our local conditions than the ;

=

SEA FISHERIES LABORATORY. 147

complete bouchot system would be—but to aid them, to
rescue from destruction the minute mussel fry in order
that they may be later on stripped off and bedded out.

The number of our mussel beds might so be largely
increased, and it ought to be re:;nembered that these
mussel beds are not valuable merely because of the mussels
they produce, but also because of the effect which they
have upon other fisheries by enriching the waters with
numerous larval and other minute forms of life. Mussel
beds become centres of attraction to numerous other
invertebrates producing swarms of living things which
are most valuable as food for young fishes, as well as for
the mussels on the bed.

th COCKLES:

529 cockles (Cardium edule) have been examined in the
Laboratory during the past year in order to find out the
food and times of spawning of this important shell-fish.

As was stated in the first report, (1892) the stomachs of
a very large proportion of the individuals are always empty,
and this seems to be the case throughout the year. It is
difficult to give an opinion as to the reason for this, as it
can scarcely be due to digestion after capture, seeing that
mussels collected at the same time and sent to the Labor-
atory along with the cockles frequently contain food.
Possibly a further examination of the cockles made at the
moment of collecting may throw some light on the matter;
it will be kept in view during the ensuing year.

In regard to the spawning time of the cockle it is diffi-
cult to give a definite period as we find them sexually
mature nearly all the year round. ‘Towards the latter
part of the year measurements were made of the ovarian
ova; and the inside of the shell and mantle cavity were
carefully washed out before opening the animal so as to

148 TRANSACTIONS LIVERPOOL BIOLOGICAL SOUIETY.

find out whether or not there were any ova free in the
cavity between the mantle and the body. ‘The sizes of
the ova are given in the accompanying lists; we usually
found a few of the ova free in the mantle cavity when the
specimen under examination was a female and when the
ovarian ova were at or about maturity.

' [ 6 female, nearly ripe. '
Grange over Sands Feb. 7 | 12 fish.|1-13 4, |@ male! nearly . He 12 Empty.

Bence wpe ANOS IO) 4s ae 14-13 ,, S a { immature 12 Empty.

ae ae Apr. 10/11 ,, |g-1 ” ; ee pone 11 Spores, ete.

Morecambe. Pooh IE TO es ee Ie ee RRO TICE. pen

Baicliff Sands. | Apr.17|23 ,, |1-1@ ,, |q2 Rmale| abouts | 93 Empty.

Morecambe. Avr. Si ala ye alee : eee ape | 12 Empty.

Eytham, Howe | apr. i8|8 4, |2-1. » |p omue| so ee oe ; 4

ee Apr. 25/12 ,, [1-14 » | dmalesperms moving] tableideNieaall

tasowe ore. Jay 9]28 ., |1-18 5, |S Rue e Te] ea Yaga
debris.

8 female, ova ‘1 m.
Duddon Sands. May 7|14 ,, |14-12 ,, |6 male, sperms not | 14 Empty.
moving.

Lyth Horse 10 female, ova ‘1 m.
ra iets Maye 9) los ot Ae 5 male, sperms not | 15 Empty.

moving.

7 female, ova ‘09-1m. 12 Vegetable

h Horse
eee” a May 23/12 ,, |1 39 "eee sperms not | Gopig
7 female, ova ‘1 m.
pythem 1 , 12 Venctaile
Salthouse Danie | ayo age ae ” eee hot | debris. .
4 female, ova ‘1 m.
Grange over Sands} May 21/12 ,, |1-1% , |8 male, sperms not | 12 Sand only.

moving.

SEA FISHERIES LABORATORY. 149

6 female ova *1-'14 m.
6 male, sperms not | 12 Empty.
moving.

4 female, ova 14 m.

Rusebeck Sands. | May 28/12 ,, {1-14 ,, |
6 male sperms moving.

Grange over Sands Meee 10 4; | 1-1k

Humphrey Head. 10 Empty.

8 female ova °15-'17m.

Lytham. May 30)12 ,, |13-14 ,, |4 male, sperms not | 12 Empty.
moving.
van 5 female, ova *15 m,
ae Cockle Jume 6/12 ,,. | 14 », |7@ male, sperms not | 12 Empty.
. moving.

Grange over Sands 4 female, ova "14 m.

Bardsea Bank. \2umell|12 ., | 1g ,, |8 male, syerms not | 12 Empty.
i moving.
; 2 female, ova "14 m.
~ ee leita A | t2 55 -| 1s », | 10 male, sperms not | 12 Empty.
moving.

Grange over Sands

‘ 6 female, mature.
Kent’s Bank. July WOj12 , ;1-1% , 12 Empty.

6 male, sperms lively.

1 female, ova ‘12 m.

7 male sperms moving
oe Bank, Sato s2 | 2-12 4, | 11 male, sperms not. 32 Empty.
moving.

13 2? immature.

Crosby ema. July 23/12 ,, | 14-14 ,, | 12 spent. 12 Empty.
7 aa uly 25} 9 $ 1-1z 4, |9 spent. 9 Empty.
ee ity 25\ ,, |1-4 ,, |.) ea 12 Empty.
Be fay 20|12 , frase, [Septet | ao mpeg
marie x22, Joan, [1omeinom tt 2) 1s pape

Grange over Sands
Grange Banks.

Duddon Cockle
Beds. ane. 14)12 ,,

9 female, ova ‘15m
5) =e) , 2g
Aug. LAT Pfam Ser 3 male sperms moving. 12 Empty.

rid 9 female, ova ‘15m.
> | 3 male sperms moving.

12 Empty.

amale r “15
Grange Banks. wept, 1d) 12>, °|/b=12' .;, : oem ALES oles Empty.

Grange over Sands} « p
Kent’s Bank. Boyes) 12,

2 female, ova *14 m. oT
a1 , 3, 10 spent. 12 Empty.

150 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Grange over Sands] Oct. 8/12 ,, |14-14 ,, aa ete ove eas Empty.

Grange over Sands

t 3 female, ova 14 m.
ie! Kent's Bank. | 0% 16/42 5, |1-18 5, :

9 spent. LO:EEp ae .

4 female, ova "165 m.

| Grange over Sands; Oct. 22}12 ,, )1-14 ,, |8 male, sperms not | 12 Empty.
i" moving.
|
Grange over Sands| Oct. 29|12 ,, |1-1% ,, | 12 spent. | _ | 12 Empty.
Grange over Sands 2 female, ova ‘15-18m| _.

Waded Scar. Nov. 5) 12 ., is ”? 10 spent. a Empty.
| 2 females, ova *14 m. | .
fe Duddon Sands, | Nov. 6/12 ,, |1-14 ,, | 4male. sperms moving| 12 Empty,
ie

ia 6 spent.

Grange Banks. Dec. 1|12 ,, | 14-14 ,, | 12 males, immature. | 12 Empty.

Kent’s Bank,

1 female, ova ‘14 m.
Cockle Beds. Deer 4) 14 5) | be

13 male, immature. 14 Empty.

2 female, ova ‘14 m.
» |1-123 ,, | 5male, sperms moving] 10 Empty.
3 male, immature.

Duddon Sands. Dec. 11 | 10

11 male, sperms mov- i
Grange Banks. Dee, 11s) 125050 a es ss ing. 12 Empty: ae

11 male, immature.

8 female ova ‘12-'15m.

Grange Banks. Dee Non Wo oe) bok :
2 9 male; immature.

‘Bet

2?

12 Empty.

In addition to the great value of cockles for their own
sake, we desire to point out that they are probably of great —
economic importance in connection with the nourishment __
of young true fish. The inter-dependence of different
forms of life in the sea is a very marked thing, the presence
of one species frequently attracts another, and sands that
contain cockles seem usually to have other burrowing —
mollusca and many amphipods and other invertebrates
and so no doubt form an attractive feeding ground for
fish.°

SHRIMPS AND SHANKS.
During the year 290 shrimps (Crangon vulgaris) have
been examined in order to ascertain further particulars in
regard to the food and times of spawning.

4
|

SEA FISHERIES LABORATORY. 151

The examination of the food confirms what has already
been stated in previous reports, that Mollusca, Crustacea,
Annelida, etc., are used as food by the shrimp. In the
stomach of one specimen examined in October, 1894,
which was one of twenty-four collected on Burbo Bank, a
perfect specimen of a young Donazx was found which
measured 1°3 millimeters, both valves being intact. The
other shrimps in this gathering had also been feeding on
the young of Donaz, but the shells were all more or less
damaged.

In regard to the spawning periods we have as yet noth-
ing further to add to what has already been stated in the
first Report, p. 34, viz., that there seem to be two chief
periods in the year, the one about November and the
other in April and May, although many are found spawning
between these periods, as the following statistics show :—

In December most of the Shrimps examined had spawn.

In February most of the Shrimps examined had spawn.

In March 60 % of the Shrimps examined had spawn. —

In April 70 to 85 % of the Shrimps examined had spawn.

In May 60 to 80 % of the Shrimps examined had spawn.

Only 20 Shanks (Pandalus annulicornis) were examined
during the year so that we have nothing further to add on
that head to what is contained in last year’s Report.

SHRIMPING AND THE DESTRUCTION OF YOUNG FISH.

We desire, however, before leaving the subject of shrimps
to bring once more prominently before the Committee, and
to recommend for careful consideration, a matter we have
discussed in a former report, (Rep. I. p. 122) viz., the
appalling destruction of young food fishes which is caused
by shrimping as practised in this neighbourhood.

In the Report for 1893 the subject was drawn attention
to, the numbers taken in particular hauls were given, and

152 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

suggestions were made as to remedial measures. The
destruction by shrimping was again referred to in last
year’s report where a full description was given of the —
“casier”’ or shrimp trap and its method of use at Le
Croisic on the Brittany coast. This instrument has not
yet had a satisfactory trial in our estuaries, and it is a
question whether even if it were shown to work as well
here as 1t does in France our men could be induced to
make use of it.

The statistics of hauls taken during the past year from
the steamer show once more, if any showing is still needed,
that that destructive engine the shrimp trawl brings up
along with a miserably small number of shrimps, an
astonishingly large number of young food fishes. On
November 2nd, off the Ribble estuary, with 5 quarts of
shrimps were taken over 5,000 undersized food fishes.
On the same date, off Blackpool with 14 quarts of shrimps
were 10,000 fish; on October 24th, in Heysham Lake
with 2 quarts of shrimps were 4,000 plaice about 4 inches
long; and so on. Of course it is satisfactory to know
that there are so many young fish on the ground, but it

is deplorable that for the sake of a quart or two of shrimps

several thousands of young fish should run some risk of
being sacrificed.

Bar SHANK NET EXPERIMENTS.

These experiments were carried out by Mr. Dawson
for the purpose of determining whether the destruction of
small fish caught while shrimping could be decreased
without affecting the number of shrimps taken. In carry-
ing out the experiments an ordinary shank net and a shrimp
trawl were worked over the same ground, along with a —
modified shank net having a bar fixed to the frame about
3 inches off the bottom, to which bar the lower part of

SHEA FISHERIES LABORATORY. 15s:

the net is attached, the three being worked simultaneously
so that the experiments might have a fair trial. As stated
in the introduction these experiments have, so far, sup-
ported Mr. Dawson’s idea as to the fish caught in this
net being fewer in number than those caught either by the
ordinary shank net or the shrimp trawl, but it would
perhaps be better that the experiments should be carried
on for a further period before any definite opinion is
expressed on this matter.

Although the results have already been given in part
in one of Mr. Dawson’s Quarterly Reports (January, 1894)
to the Committee, we consider that their importance
renders it desirable that they should have if possible a
more extended circulation and so we state them here more
fully taking the more striking examples for the various
months.

(1) In December, 1893, an experiment was made on
the Blackpool closed ground with a Bar Shank net, an
ordinary shank net, and a shrimp trawl, the three being
worked simultaneously, with the following result :—

The Bar Shank net caught 3708 fish and 11 quarts of

Shrimps.

The Ordinary Shank net caught 5221 fish and 104

quarts of shrimps.

The Ordinary Shrimp trawl caught 20889 fish and 223

quarts of shrimps.

(2) In January, 1894 another experiment was made on
the same ground and under the same conditions with the
following results :—

Bar Shank net, 2011 fish and 14} qts. shrimps.

Ordinary Shank net, 2027 fish and 8} qts. shrimps.

Ordinary Shrimp trawl, 8512 fish and 6 qts. shrimps.

(3) An experiment in the Ribble carried out under the
same conditions in January gave the following :—

= nee on en
Bib!
ey

‘154 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Bar Shank net caught 746 fish and 114 qts. of shrimps.

Ordinary Shank net, 561 fish and 8} qts. shrimps.

Ordinary Shrimp trawl, 1950 fish and 6 qts. shrimps.

(4) Another experiment was made on the Blackpool
ground with the three nets in the month of March and
the following is the result :—

Bar Shank net, 38 fish and 13 qts. shrimps.

Ordinary shank net, 865 fish and 9 qts. shrimps.

Ordinary Shrimp trawl, 3076 fish and 23 qts. shrimps.

(5) In the same month (March) an experiment was
made in the Mersey with the three nets with a result as -
follows :—

Bar Shank net caught 536 fish and 12 qts. shrimps.

Ordinary shank net, 365 fish and 63 qts. shrimps.

Ordinary Shrimp trawl, 1177 fish and 13 qts. shrimps.

(6) In August another experiment was made on the
Blackpool closed ground with the following result :—

Bar Shank net caught 502 fish and 2 qts. # gill shrimps.

Ordinary Shank net, 330 fish and 2 qts. shrimps. |

Ordinary Shrimp trawl, 3362 fish and 2 qts. shrimps.

‘Drift BotTTLES AND SURFACE CURRENTS.”

In connection with the investigation of the surface life,
in discussing the appearance and disappearance of swarms
of certain Copepoda and Meduse, and in considering the
possible influence of the movements of such food matters
upon the migrations of fishes, and also in connection with

the movements of the fish ova and floating embryos, 1t
occurred to us, that it would be worth while to try to

ascertain the set of the chief currents, tidal* or otherwise

* The tidal currents of the district are already to some extent known, and
are marked in the charts and given in books of sailing directions, as Admiral ~
Beechy’s ‘‘ Tidal Streams of the Irish Sea ;” but we desire to ascertain the
resultant currents from all influences which would affect the drift of small —
floating bodies.

SEA FISHERIES LABORATORY. 155

such as the movement of surface waters caused by preva-
lent winds. The Prince of Monaco started a few years
ago the system of distributing over the North Atlantic
large numbers of small floating copper vessels, with the
object of finding out where they drifted to. This plan we
have adopted, with slight modifications, and in September
we started the distribution of what may be called “ drift
bottles” over the Irish Sea. A small, strong, buoyant
bottle, measuring 7°5 cm. by 1°8 cm., which seemed well
_ suited for the purpose, and which costs only 7s. per gross
was selected. A notice was drawn up, as follows, to goin
the bottles, and a large number of copies were printed and
numbered consecutively.
Any one who finds this is earnestly requested

to write the place, and date when found, in the

space (on the other side) for the purpose, place

the paper in an envelope, and post it to

PROFESSOR HERDMAN,
University College,

LIVERPOOL.
TO ores ose Ss

Postage need not be prepaid.
Turn over.
[OTHER SiDE.]
Please write distinctly, and give full particulars.
MMMM. WACEC. TOUNG sys acme de nvaduia ces dssavevsesds

2 7

ere eee eee eee eee eee eee eee eee eee eee eee eee eee eee eee eee eee eeeeeeeee

DATE, when found
Name and address of sender.

ee 2

eer eee eee eee eee eee eee ees eeeeseeeeeee eee eeeseseeesesese

A paper was then placed in each bottle, so folded that
the number could be readily seen through the glass, the

156 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

cork was well pressed down, and dipped in melted paraffin.
Over a hundred of these bottles have, since September
80th, been dropped into the sea in various parts of our
area, a record being kept of the locality and time when
each was set free. Several dozen were let off from the
Isle of Man steamer in crossing to Douglas and back, at
intervals of quarter of an hour, and from our trawler when
dredging between Port Erin and Ireland. Several dozen
have been let off from Mr. Alfred Holt’s steamers in going
round to Holyhead and in coming down from Greenock.
Mr. Dawson on the fishery steamer ‘“‘ John Fell”’ has dis-
tributed a number along the coast in the northern part of
the district, and others have been set free at stated inter-
vals during the rise and fall of the tide from the Morecambe
Bay Light Vessel, and Lieutenant Sweny has kindly
arranged to have a similar periodic distribution from the
Liverpool NorthWest LightVessel. Altogether, over 33 per
cent., or about one in three of the papers distributed have
been subsequently picked up on the shore and returned
duly filled in and signed. They come from various parts
of the coast of the Irish Sea—Scotland, England, Wales,
Isle of Man, and Iveland. Some of the bottles have gone
quite a short distance, having evidently been taken straight
ashore by the rising tide. Others have been carried an
unexpected length, e.g., one (No. 35), set free near the
Crosby Light Vessel, off Liverpool, at 12.30 p.m., on
October 1st, was picked up at Saltcoats in Ayrshire, on
November 7th, having travelled a distance of at least 180
miles* in 87 days; another (H. 20) was set free near the
Skerries, Anglesey, on October 6th, and was picked up

one mile N. of Ardrossan, on November 7th, having

travelled 150 miles in 31 days; and bottle No. 1, set free

*More probably, very much further, as during that time it would
certainly be carried backwards and forwards by the tide,

a

;

SHA FISHERIES LABORATORY. 157

at the Liverpool Bar, on September 30th, was picked
up at Shiskin, Arran, about 165 miles off, on November
12th. On the other hand, a bottle (J. F. 34), set free on
November 7th, at the Ribble Estuary, was picked up on
November 12th, at St. Anne’s, having gone only 4 miles.

It would be premature as yet—until many more dozens
or hundreds have been distributed and returned—to draw
any very definite conclusions. It is only by the evidence
of large numbers that the vitiating effect of exceptional
circumstances, such as an unusual gale, can be eliminated.
Prevailing winds, on the other hand, such as would usually
affect the drift of surface organisms are amongst the nor-
mally acting causes which we are trying to ascertain. We
may, however, state for what they are worth, the following
results obtained so far :—(1) Nearly 50 per cent. of the
bottles found have been carried across to Ireland, and
they are chiefly ones that had been set free in the southern
part of the district (between Liverpool and Holyhead) and
off the Isle of Man; (2) the bottles set free along the
Lancashire coast and in Morecambe Bay seem chiefly to
have been carried to the South and West—to about
Mostyn and Douglas; (8) it is apparently only a few that
have been carried out of the district through the North
Channel. Perhaps, the most interesting point so far is
that so many of the bottles have been stranded on the
Trish coast, although they were sent off for the most part
much nearer to the Enghsh and Welsh coasts, showing
probably the influence on the tidal currents of the spell of
Easterly winds in October. It is interesting to learn that
the Fishery Board for Scotland has also commenced a
similar inquiry to ours by the distribution of floating
bottles in the Scottish territorial waters. No account of
their experiment has yet appeared, but it will be of some
importance to compare results with them, say at the end
of the first year’s work.

he
r
\
.

' i A ”
158 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

_Faunistic INVESTIGATIONS.
sepumans carrying out the work of examining ane fish |
caught in the net while on board the steamer a sharp
look-out is kept upon the debris brought up along with

_ the fish so that we may know as much as possible of the

nature of the food at the bottom, and so that any rare
vertebrates or invertebrates that may happen to be among
the refuse may not escape notice. It is only by having
a thorough knowledge of the fauna of the district that we
ean expect to be able to make out accurately the food of
the various fishes, for without such a detailed knowledge
of the species one would often be puzzled to tell what the
fish really are feeding upon, the contents of the stomachs
get so much altered during the process of digestion.

As a result of the untiring zeal of the various members
of the Liverpool Marine Biology Committee and other
workers it is now no easy matter to make many additions
to the fauna of the district, the records during the past
year being only 1 Fish, 1 species of Cumacean, 5 species
of Copepoda, 1 Ostracod and 1 Sponge.

The following are the names of the various species

referred to, with a note to each of some particulars which

may be of interest.

FIs.

Zeugopterus unvmaculatus, (Risso).
1880-84, Day, Brit. Fish., v. I1., p. 17, pl. xcix., fig. 1.
1885, Brook, Appendix to Fourth Ann. Rept. Fish.
Board for Scotland, p. 225, pl. 1x. :
Four specimens of this rare and pretty little fish were
captured by the trawl-net of the Fisheries Steamer “ John —
Fell,” while trawling on the off-shore grounds 10 to 12 —

- miles west from Morecambe Bay Light Vessel during the

month of May, 1894.

SEA FISHERIES LABORATORY. 159

Depth 23 fathoms, bottom sand, shells, and stones.

The specimens agree in all points with the figure and
description given by the late George Brook in the Scottish
Fishery Board’s Fourth Annual Report. The figure given
in Day’s British Fishes does not seem to be very charac-
teristic.

One of the fish was a mature female and measured 54
inches in length, the unfertilised ova measured 1:078
millimeters and the oil globule ‘1848 millimeter.

CUMACEA.

Petalosarsia declivis (G. O. Sars).
Petalomera declwis, T. Scott, Eleventh Annual Rep.
Fishery Board for Scot., Pt. IIT., p. 215, pl. v., fig. 43.
A few specimens of this Cumacean were obtained among
bottom tow-net material collected 14 miles N. by W. from
Liverpool N.W. Light Vessel. The tow-net had touched
the bottom and brought up a small quantity of sandy
mud which was carefully washed through a muslin sieve
and on examining the contents of the sieve this species
was found, and also a copepod which is referred to later
on. This species had not previously been recorded for
the west coast of Britain.

COPEPODA.

Longipedia minor, T. and A. Scott.

Longuypedia coronata, var. minor, T. and A. Scott.
Hleventh Annual Report, Fishery Board for Scotland,
Part ILI, p. 200, pl. 11., figs. 14—-20.

In rock-pools on Hilbre Island, collected by hand net,
not common. ‘This is one of the copepods we sometimes
find in the stomachs of young fish from the district.

Bradya minor, T. and A. Scott.

Several specimens of this new Bradya were obtained
in the same gathering with the last.

160 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Lictinosoma norman, T. and A. Scott.

Amongst material collected in Barrow Channel by
Professor Herdman while surveying the ground for the
proposed establishment of Mussel Beds, &c., in the vicinity
of Piel and Roe Islands.

Pseudanthessius sauvage, Canu.

Pseudanthessius sauvaget, Tl. Scott, Twelfth Apainted
Report, Fishery Board for Scotland, p. 260.

Washed. from the Echinoderms, Spatangus purpureus
which were brought up in the trawl net of the steamer
while working on the off-shore grounds during the year.
It has also been found on Hchinocardiwm cordatum, see
Canu’s monograph ‘“‘ Les Copepodes du Boulonnais.”

Lichomolgus agilis heydig).

Lichomolgus concinnus, T. Scott, Tenth Annual
Report, Fishery Board for Scotland, Part a p. 261,
pl. xi., figs. 25—33. -

A few specimens were found amongst the bottom a

material from which Petalosarsia were obtained.
This is not the Lichomolgus agilis of T. and A. Scott

_ referred to in the First Report as having been found in

the cockles. The latter species is identical with Herman-
ella rostrata, Canu, a species which was described and
published a short time before the figures and description
by T. and A. Scott appeared in the Annals and Magazine

of Natural History for September, 1892.

OSTRACODA.

Cytheropteron pyramidale, G. O. Sars.
Among the mud brought up in the tow-net gathering
in which Petalosarsia was found. . :

' SPONGE.

Lewosella (Spongionella) pulchella, Sowerby.

SEA FISHERIES LABORATORY. 161

Amongst trawl refuse, which had been collected 14 miles
N. by W. from Liverpool N.W. Light Vessel in the same
haul from which Petalosarsia was got in the tow-net.
This sponge had not been taken before on the west coast
oi Britain, and we are indebted to Dr. Hanitsch for the
name.

Nore. It may be of interest to record here the occur-
rence of the Calanid Candace pectinata, Brady, in the
Trish Sea. It was found among some surface tow-net
material collected by Mr. R. L. Ascroft off Lambay Island
on the Irish coast, about 10 miles N.E. of Dublin, while
on board a trawler working there in November, 1894.
Mr. Ascroft also kindly sent to the Fishery Laboratory
for examination some material which he obtained when
trawling off Galley Head, Co. Cork. This has been
examined by Mr. A. O. Walker who reports the following
interesting and rare Amphipods and other higher Crustacea,
viz.:—Henulamprops assimilis, Diastylis (? n. sp.), Ciro-
lana borealis, Parathenusto oblivia, Callisoma crenata,
Hiuppomedon denticulatus, Orchomenella ciliata, Typho-
sites longupes, Lepidepecreum carimatum, Ampelisca spini-
pes, Monoculodes carinatus, Hpimeria cormgera, and
Paratylus vedlomensis.

SUMMARY AND CONCLUSIONS.

In concluding this Third Report on the work of the
Fishery Laboratory we desire to point out that there are
several matters to which we have drawn attention in this
and in former reports and which seem to us still to be
pressing for attention and to be well worthy of serious
consideration. ‘hese are :—

1° In regard to Shrimping :—A fair trial of the Croisic
trap (see Report II., p. 128) and a trial of artificial Shrimp
culture in enclosed areas (Report I., p. 184) with the view

162 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of providing some substitute for the present destructive
method of fishing.

_ 2° The erection of some a Mussel bouchots so as ‘o
catch spat (Report II., p. 183) with the view of forming
fresh Mussel beds or re-stocking old ones.

3° The establishment of a Sea-fish hatchery (this Rep.,
p. 186) with the object of doing all that is possible to add
to the population of the sea.

Other points which we Baw raised in these reports,
such as the determination of the spawning grounds, feeding
grounds and nurseries, the question of the “vitality” of
fish caught in trawls, the distribution of immature fish,
the size at maturity and the benefit of the imposition of a
‘size limit,” have all now been more or less completely
determined and settled. There still remain, however,
unsolved or only partially solved problems, and new ones
are arising as a result of the discussion of our statistics,
both these collected in the laboratory and also those taken
at sea by the steamer in carrying out the scheme of
investigation recommended in our first Report.

_ We propose in the coming year, in addition to the
usual statistics on the food, maturity, growth, &c., of -
fishes, and any questions that may arise therefrom, to
investigate more closely the shell-fish on the beds, and to
pay special attention to the spawning of fish and to the
larval and other immature stages.

163

[WORK FROM PORT ERIN BIOLOGICAL STATION, ]

Note upon the yellow variety of Sarcodictyon catenata,
Forbes, with remarks upon the GENUS and its species.

By W. A. Herpman, D.Sc., F.B.S.,

PROFESSOR OF NATURAL HISTORY IN UNIVERSITY COLLEGE, LIVERPOOL.
[Read December 14th, 1894.]
Plate VITI.

In 1883, I published a somewhat detailed account* of the
structure of the ordinary form of Sarcodictyon catenata,
Forbes, from the examination of specimens dredged at
Lamlash and in Loch Fyne, on the West Coast of Scot-
land, in 1880 and following summers. Some of my
colonies from Loch Fyne were of a pale yellow colour,
and an investigation of these showed that they only
differed from the red colonies in having the numerous
calcareous spicules of the mesogloea perfectly colowrless
in place of being of a dull red tint. Hence, I came to the
conclusion that here, just as in Alcyonium digitatwm,
where several distinct colours of colony are found, the red
and the pale yellow colonies were merely cases of colour
variation, and that all the specimens that I had examined
were to be referred to the one species, Sarcodictyon
catenata. Moreover, I made the suggestion that probably
Forbes’ second species, S. agglomerata, which was said+
to differ from S. catenata in being of an “‘ ochraceous
colour, and in having the polypes placed in little

>

yellow ’
eroups of from three to five instead of in single file,
would prove to be the same species, since (1) several of
my red colonies had the polypes in groups, and (2) the
yellow colonies bridged the gap as regards colour.

* Proc. R. Physical Soc. Edinburgh, vol. viii., p. 31.
+ Trans. Roy. Soc, Edinburgh, vol. xx., p. 307, 1851.

164 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Three years later, in 1886, I added* some further
~ details in regard to Sarcodictyon catenata in the living |
condition from colonies dredged off Port Erin, in the Isle
of Man, and which I had been able to keep alive for some
time. In that paper I figured the polypes in the fully
expanded condition, showing that the polype may extend
to three times its usual height (7.e., to 5 or 6 mm.), the
translucent white upper expanded part of the body being
about twice the length of the lower opaque red-coloured
part. The figures (loc. cit. pl. 11.) showed a colony of
thirteen polypes fully expanded, natural size, and one
expanded polype, magnified, like the one now reproduced

here (PI. VIIL., fig. 3).

Since that date we have frequently, during the
L.M.B.C. expeditions, dredged colonies of the red
Sarcodictyon catenata off the south and west of the Isle
of Man, at depths of from 10 to 40 fathoms, and generally
attached to shells or to stones. ast summer (1894) in a
haul of the dredge taken at 2 miles off the north side of
the Calf Island on August 25th, depth 22 fathoms, bottom
sand and shells, I was fortunate enough to obtain several
colonies of yellowish tints, which I think are quite —
sufficient to show conclusively that Forbes’ Sarcodictyon
agglomerata is the same species as S. catenata. I had
previously shown that there were red colonies in the
agglomerated condition, and yellow colonies with the
polypes in single file. Now I have yellow colonies of
several tints in the agglomerated state, and one of these,
in its ‘‘ ochraceous yellow ”’ colour and in the grouping of
polypes in threes and fives, exactly corresponds with
Forbes’ original description (see Pl. VIIL., fig. 1).

On cutting sections of some of these yellow colonies I
find that they have not all colourless spicules only, as I

* Fauna of Liverpool Bay, vol. i. p. 120.

NOTE ON SARCODICTYON. 165

found was the case with the Loch Fyne specimens in 1883,
but that there are two distinct tints of yellow colony, one
paler and greyer (Pl. VIII., fig. 2) in which the spicules
are all colourless and the yellowish tint is merely due to
the superficial tissues (chiefly the ectoderm) of the polypes
and stolon, and the other of a much richer yellow (PI.
VIIL., fig. 1), with sometimes a tinge of orange, and in
these most of the spicules are very distinctly of a yellow
colour, even when seen singly in thin sections (PI. VIII.,
figs. 6, 7, 8), and when in mass along with the soft
tissues they make up the rich yellow seen on the outside
of the colony. The spicules which are yellow in these
cases are exactly the ones which are red in the red
colonies, viz., those of the stolon and of the lower thick
opaque part of the body-wall, leaving those of the upper
translucent part of the body-wall and of the tentacles
colourless (see Pl. VIII., fig. 7). I succeeded in keeping
yellow colonies, of both tints, alive and fully expanded in
the aquarium at Port Erin for some time last summer.
The upper parts of the body wall and the tentacles are
exactly like those of the red form (see Pl. VIILI., figs. 4, 5).

Consequently, there are altogether three colour varieties
of Sarcodictyon which I know of :—

1. The red form, with red spicules in the stolon and
lower part of the polype;

2. The bright yellow form, with yellow spicules in
the stolon and lower part of the polype;

3. The pale yellow form, with colourless spicules
throughout the colony.

All of these usually have the polypes in single file, but
any of them may exceptionally have the polypes grouped
in little clumps (as in Pl. VIII, fig. 1). Consequently,
the characters distinguishing Sarcodictyon agglomerata

Ae Pe eee hike meee te ee ee aia oe

“ a » ’ at vs Pe oe

166 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

fall to the ground, and that name must be regarded as a
synonym of S. catenata, Forbes.

_ In all the other details of structure as seen in working
through thin sections I find that the yellow colonies agree
with the ordinary red ones—the only difference is in the
colour of the spicules. I have been fortunate in coming
upon ova in the yellow colonies recently examined—I
had failed to find any trace of reproductive organs in the
specimens examined in 1883. The ova are of rather large
size, 0°09 mm. in diameter, and have a very distinct
germinal vesicle and spot. They are enclosed in cellular
follicles, about 0°12 mm. in diameter, which hang by long
narrow pedicles from the edge of the mesenteries—project-
ing into the inter-mesenteric cavities in groups of two
three and four together (Pl. VIII., fig. 10). I have seen
no trace of spermatozoa, so it is very possible that the
colonies are unisexual and that I have only examined
female colonies.

In conclusion, I desire to make a few remarks in regard
to the genus. Prof. Hickson has recently published a,
| paper* on the genera of the Alcyonaria Stolonifera in
which he deals critically with the various genera which
have been placed in the family Clavularude, and in the
course of his analysis of this group he refers (loc. cvt. p. 832)
briefly to Sarcadictyon and comes to the conclusion that
it must be merged in the genus Clavularia. There is no
doubt that the three genera Sarcodictyon, Rhizoxenia and
Clavularia are, if distinct, very closely related groups,
and our British species Sarcodictyon catenata has by some
writers been placed in Rhizorenia and by others in Clavu- :
laria. It cannot, however, be referred to Rhizoxenia as
that eae was characterised by Ehrenberg, its founder, and

“* Trans, Zool, Soc. Lond., vol. XIII, pt. IX, p. 325, Oct. 1894,

NOTE ON SARCODICTYON. 167

since that by Milne-Edwards and Haime, as having the
polypes non-retractile, while S. catenata has eminently
retractile polypes.

There remains then the question, ought our species to
be included in Clavularia ? If it cannot be separated
from the species of that genus, Clavularia as a name has
priority, having been established by Quoy and Gaimard in
1834 while Sarcodictyon catenata was not discovered by
Forbes till 1845 and was not described till 1847. I am of
opinion, however, that the constitution of the polype wall
out of two very distinct parts, a lower thicker opaque
coloured part into which the upper thinner translucent
colourless part can be retracted, is so different from the
condition seen in the species of Clavularia (see Hickson’s
beautiful plates, which show, in all the species figured,
the body-wall extending in an unmodified state up to the
bases of the tentacles) that 1t gives us sufficient grounds
for the separation of S. catenata generically. And although
the character of the body-wall above referred to was not
included in the original description of Sarcodictyon, still
the species S. catenata, Forbes, was the type of that
genus and consequently Sarcodictyon is the name that
ought to be retained as the generic title.

As to the further question whether there are any other
species of Sarcodictyon, that is a very doubtful matter.
Hickson mentions two species in all,—the well-known
S. catenata, and another ‘‘S. colinabum”’ which I have
never heard of. He gives as the locality Scotland, and
as authority ‘‘ Forbes (?),” but I cannot find the name in
Forbes’ works, and am inclined to think there must be
some mistake about it.* Hickson does not include in his
list of the species he desires to add to Clavularia, Sarco-

* Since the above was printed I have heard from Prof. Hickson that he

possibly mistook a note book entry of S. agglomerata for **S. colinabwm.”

a : P ats -
fos

168 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

dictyon agglomerata, Forbes, and S. rwgoswm, Pourtales.
I have already expressed my opinion above that the former __
of these must be merged in S. catenata, and I feel that we |
require further detailed information about Pourtales’ West
Indian form before being able to decide either that it is
distinct from 8S. catenata or that it really belongs to the
genus Sarcodictyon.

EXPLANATION OF PuatTE VIII.

Fig. 1. Colony of bright yellow form of Sarcodictyon
catenata, (this is the S. agglomerata of Forbes), —
nat. size, painted from life.

. Colony of pale yellow form, nat. size.

. Expanded polype of ordinary red form, enlarged.

. Expanded polype of yellow form, enlarged.

. Anterior end of another expanded yellow polype.

Es)

ai
fee Ua
Oo Oo - W LO

. Series of spicules from the body-wall of a yellams

form (xX 200). ;
Fig. 7. Part of a horizontal section through a yellow

colony, showing two polypes cut in section, with

yellow spicules in the outer body-wall andin the _

stolon, and colourless spicules in the invaginated

part of the polype (x 50). :

Fig. 8. Part of a section of a yellow polype showing zone

of spicules (X 50). Ee,

Fig. 9. A vertical section of a pale yellow polype (x 50). |

Fig. 10. Part of a transverse section of a yellow pol e

i

showing the ova in their capsules attached — a2
pedicles to a mesentery, (X 200).

169

[WORK FROM THE PORT ERIN BIOLOGICAL STATION. |

OBSERVATIONS on the TUBE-FORMING HABITS
of Panthalis oerstedt.

By Arnotp T. Watson, Sheffield.

With Plates IX.—xX.
[Read January 11th, 1895.]

On the 5th of June, 1893, when dredging with Professor
Herdman and the Members of the Liverpool Marine
Biology Committee on board the s.s. ‘‘ Mallard,” at a
depth of 60 fathoms, 14 miles West of Dalby (Isle of Man)
a number of long, curious, soft muddy masses were brought
to the surface, which in the absence of knowledge of their
character were promptly, and for the time being, appro-
priately described as ‘‘mud sausages.” These masses
were at once carefully examined with the object of ascer-
taining whether they contained any tenant. The search
was unsuccessful, except as regards one mass, from which,
after long and careful manipulation under water, I
succeeded in expelling a living specimen of Panthalis
oerstedt, about 2 inches long.

Since that time, a fair number of these mud masses
(which on examination proved to be mud tubes) have
been brought up when dredging in deep water off the
Isle of Man. Many of them were empty, but others
inhabited; the most frequent tenant being Panthalis ;
and the question arose whether Panthalis was the
fabricator of the tubes, or simply a tenant; andif the
former, by what method such curious heavy structures
were made.

Panthalis is a Polynoid worm of the sub-family Acoé-
tidee, all the members of which are rare. A few of them,
dwellers in southern waters (Polyodontes mazillosus,

170 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Acoétes pleet, Huarche tubsfex, and perhaps others) are
recorded as ‘‘ dwellers in tubes;”’ but any such habit on
the part of the northern form, Panthalis, appears to have
been unknown, and Kinberg, who describes and figures
the animal, makes no reference to it.

From the fact that Panthalis when found off Port
Erin has always been associated with these mud tubes,
it seemed most probable that it was the maker of them:
still, this could only be accepted as a probability, and it
was obviously most desirable, that if possible, a8 question
should be definitely settled.

The tube-masses, which are very soft, and easily pulled
asunder, are usually about 14° to 13” in external diameter
by about 3} inches long, with loose mucus-like extensions
at either end, thus concealing the entrances. The
estimated internal diameter of the tubes is usually about
3”. Careful examination, by sectionising, shews the
thickness of the walls in the centre to be about +”; these
are composed of a number of thin layers; the one nearest
the animal being simply a coating of mucus-lke threads,
slightly sprinkled over with fine mud, followed by other
thin layers, distinguished by colours varying from dark to
reddish-brown, and consisting of a felty structure in which
mud is loosely entangled, until the outermost layer is
reached, which is generally of a drab colour, and very loose
in texture. In the early part of last year I embedded
portions of these tubes in paraffin, and prepared others by
staining, hardening and grinding-down, to form thin
sections, with the object of ascertaining, if possible, the
method of fabrication; but these preparations, though
now valuable, at first afforded very little information, and
I was consequently wishful to observe the habits of the ©
living worm, an opportunity for which was last summer
afforded me by Professor Herdman.

TUBE OF PANTHALIS. alr ¢i

On the 25th of August he kindly took me on the s:.s.
“Albatross” to the ground on which Panthalis had pre-
viously been found, and we were fortunate in securing
two promising mud masses, which, with a supply of mud
from the same spot, I took to the aquarium at Port Erin.
About noon on the following day these masses were placed
close against the glass sides, in different parts of a small
portable tank, and deposited in the aquarium building.
The water was changed by syphoning, twice daily, to
correspond with the tides (this was a needless precaution,
but I wanted to avoid any chance of failure), but no mud
was in the first instance supplied, as I wished to make the
worms, if present, shew the position of the openings of
their tubes, so that I might re-arrange them if unfavourably
placed for observation; or take means of guiding them,
if needful, in the direction of the glass. At 10.30 p.m.
one of the tubes displayed an opening at each end, but no
animal was yet visible. At 7 o’clock next morning,
however, at the opening away from the light, the tip of
the head, and two long palps were visible, but the animal
was exceedingly sensitive to vibration, and on my approach
immediately retreated. Having ascertained my ‘“‘bearings”’
I, about noon, covered the floor of the tank, round the
tube, with some of the fine mud brought for that purpose,
but restricting the depth to half an inch, in order that any
movement of the animal might be traced. The tank was
carefully watched, but no change took place during that
day; on examining it next morning, however, there were
signs which led us to think that the worm had, during
the night, travelled five or six inches on the surface of the
mud, and returned to its starting point, whence it had
burrowed along the front of the tank, forming a channel
the whole length between the glass and the underside of
the old tube. The latter action was all that one could

172 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

desire, and was in fact the exact state of affairs I was
aiming to produce, but had not hoped to attain so easily.
This habit of annelids in captivity burrowing or building
against the glass, has often been helpful to me. It affords
excellent opportunities for studying the form and habits of
the animals under natural conditions, the interior of the
channel being exposed to view. By carefully changing
the water (by syphoning) the worm was soon made to
feel at home, and except for the occasional vibrations to
which it was always most sensitive, seemed to be unaware
of the shallowness of the water to which it had been
transferred. At first Panthalis was somewhat cautious
about exposing itself in the clear channel, and the only
parts visible were the head, and two long palps; between
which, two red spots were seen, and on either side the
cirri of the first pair of parapodia. These were seen
peeping out into the new channel, the remainder of the
body of the worm being hidden in the old tube, or under
the mud which had been placed on the floor of the tank.
Gradually the animal gained confidence, and during
the day an inch of its length was exposed to view, and by
night almost the full length of its body was reposing
quietly in the new channel, the posterior extremity only
remaining buried in the old tube. Upon examination by
lamp-light, I found that the glass, just at the commence-
ment of the clear burrow, was for a very short distance
covered with mucus-like threads forming a ‘‘ cobweb.”
The animal itself was at first very sensitive to the artificial
hght, but subsequently bore it better, though it evidently
felt the heat, as it contracted the portion of the body on
which the hght was concentrated. As seen in its burrow,
the animal possessed 60 pairs of parapodia, and was about
83 inches long, (the tube in which it was captured being
about half an inch longer). The width, across the

TUBE OF PANTHALIS. ifs

animal’s back, to the extremities of the parapodia, was
about =; the two sides being approximately parallel,
excepting just at the two extremities, where the outline of
the body tapered off markedly.

In a dorsal view were seen the two greyish-flesh
coloured palps, extending about 3, of an inch in front of
the prostomium ; on either side of the palps, the cirri of
the first pair of feet were visible (appearing like antenne),
and apparently between the palps, two eyes, with reddish
stalks (between which is a minute tentacle) occasionally
peeped out from beneath the first pair of elytra. Except-
ing at the two extremities, where the elytra of the sides
meet and overlap, rather more than the central third of
the back was uncovered, the anterior part of its length
being of a pearly white, with a central pink stripe.
Further on it was flesh-coloured, due to the internal organs
being partially visible through the semi-transparent, finely
striated integument. ‘The dorsal blood-vessel was most
striking. For the greater length of the body it undulated
just beneath the skin, and a beautiful red, bead like effect
was produced. About one third the width of the back,
on either side, is covered by the pearly-white, semi-trans-
parent elytra, the first few pairs of which are flat, and
the remainder campanulate. During life, these do not
rest upon the body, but in front are tilted-up, so as to
meet at an angle above the prostomium, the last few pairs
of elytra also assuming a similar position. Possibly this
rule may apply to the intermediate elytra, but I had no
opportunity of ascertaining this.

A constant rising and falling of the elytra, as though
to facilitate the passage of water for purpose of respiration,
was observable, and the positions of the parapodia were
dimly indicated through them. Laterally, the anterior
portion of the animal was most interesting, as in addition

174 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

to the organs mentioned as visible in a dorsal view, the
second pair of parapodia, to which are attached special
and important functions, came into sight. These limbs
carry the first pair of elytra, and possess parts, equivalent
to those in succeeding parapodia, modified and adapted
to their work, which, as will be explained, is chiefly that
of forming the tube inhabited by the worm. Their ventral
cirri are much longer, and apparently stronger, than those
which follow; their tips are semi-transparent, but for
the greater part of their length they are of an opaque
whitish colour (other ventral cirri are in parts of a
reddish tint) and being situate near the opening of the
mouth, have the appearance of buccal tentacles, or bristles,
which are usually directed forwards and downwards, the
tips almost meeting. The parts which correspond with
the ventral and dorsal lobes of other parapodia, are
modified in shape, and opaque white at their extremities ;
but these limbs are twisted downwards on their axes, so
that the portions which correspond to the dorsal lobes in
other parapodia, instead of being above, are in advance
of the ventral lobes. They are striking objects even to
the naked eye. The sete also differ, in that they are
chiefly long and straight or regularly curved, and although
some of them may be described as weak bristles, these
are spear-like in form, and devoid of the short terminal
‘‘whip”’ or hair, which is characteristic of Panthalis.
The long curved setze, which issue from the cleft between
the lobes of the parapodia, are directed across the mouth
of the worm, and their object is probably to draw the
threads across, in the weaving process shortly to be des-
cribed. In this view, too, the outline of the front portion
of the body is noteworthy. At a distance of about 33 of
an inch from the extreme front, the dorsal integument,
which is here less covered by elytra than further on

gf _'

_ TUBE OF PANTHALIS. 175

was swollen up, giving the appearance of a shallow blister
about $ an inch long; while ventrally commencing at a
somewhat corresponding point the integument rose
abruptly, forming a sac over the blood vessel and pro-
ducing the effect of a broad longitudinal keel. This
portion of the body is capable of very considerable
expansion, the advantage of which will shortly appear.
In ventral view and outline, the sides of the body, inclu-
ding the parapodia, are approximately parallel, the anterior
extremity being bounded by a semi-circle, and the posterior
by a curve more or less acute according to the elevation
or depression of the elytra. Forward, for about ~ of an
inch, stretched the two palps, on either side of which the
cirri of the first pair of parapodia (as antennse) were
sometimes visible ; and between the palps, rising from the
under-side of the eyestalks, two small tentacles; while,
with a lens, the tip of the small median prostomial tentacle
could also be seen. About ;4, of an inch from the extreme
anterior boundary of the prostomium, are situate the two
buccal bristles or tentacles, one on either side of the
mouth, which appears as a short longitudinal line extend-
ing forwards between these tentacles. This line indicates
the depression, into which the integument folds in graceful
curves from every direction, when the proboscis is retrac-
ted. Just in front of the buccal tentacles, occupying the
space between these and the bases of the palps, come
the ventral and dorsal lobes, of the second pair of para-
podia, or as I propose calling them the ‘‘ weaving feet.”’
Centrally situate, at a distance of about 3, of an inch
behind the bases of the buccal tentacles, the dark red
ventral blood vessel was a most conspicuous feature. Its
front boundary was square and broad, about -/; of an inch
wide, but the vessel was rapidly reduced in size, and after
a course of say half an inch, ran as a moderately fine red

176 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

line to the posterior extremity. After death the course
of this vessel is marked by a longitudinal ventral ridge
which has been referred to by a previous writer as the
covering of the nerve cord. The ventral surface of the
animal was opalescent, with a golden shade towards the
sides, caused probably by the spinning glands and sete of
the parapodia. The posterior extremity of the body was
broadly forked, owing to the last pairs of a being
usually directed backwards.

As specimens in alcohol are in every way much con-
tracted and altered, this description of the large Panthalis
under observation will probably be of interest, the dimen-
tions given being a rough guide to the relative positions
of the organs, though other specimens which I have
examined vary very considerably both in size and colour.

The third day after being deposited in the tank, the
animal was evidently quite settled, and it proceeded to
complete the burrow against the glass, with the object of
making an opening at the left hand extremity of the old
tube. Harly in the morning of that day I found Panthalis
had forced about 3 of an inch of its body through the mud
at this point, and was ‘‘ prospecting ;”’ on my approach
this was quickly withdrawn; but the animal remained in
the burrow the greater part of the day, thus affording
ample opportunity for observation of which I availed
myself by making careful sketches both with and without
the microscope. The most noteworthy occurrence of the -
day was an example given by the animal of its power to
very largely expand the forepart of its body, which it
utilised for enlarging the diameter of its burrow. By
this means it lifted the old heavy tube fully ¢ of an inch,
and almost rolled it way from the glass. By night, a
clear passage and opening into the water had been made.
The following morning I again found Panthalis, lying

TUBE OF PANTHALIS. Ce

on its back, with about $ an inch of the forepart of its
body projecting from the opening most recently made.
During the night it had searched round the mouth of
the old tube for a distance of about an inch, and had
lined the new mouth of the burrow with ‘“ cobwebs.”
In the course of the following twelve hours it partially
buried this cobweb with mud which it had drawn in at
the opening, and it was evidently commencing a solid
tube. It also now shewed great activity by burrowing
under, and violently upheaving the mud at the opposite
end of the burrow, with the object of making a second
opening in that direction, which it ultimately accom-
plished. I therefore watched it until 11 p.m. and made
a careful sketch of the floor of the tank, in order to
observe any change which might take place during the
night. Just before leaving, I found that a considerable
amount of ‘‘cobwebbing”’ had been done during the
evening, and I saw the animal, for the first time, reverse
its position in the burrow, by doubling upon itself ventrally.
On returning to the aquarium at 7 o’clock next morning,
I found Panthalis, as usual, with its head just at the new
mouth of the burrow, to which, it had, during the night,
added a piece of tubing between the glass of the tank, and
the opening of the old tube; the material for which had
been obtained by digging a hole about 4 inch diameter
and + of an inch deep, in the mud close by. As no food
had yet been supplied, I now attempted to feed Panthalis
by offering it a small red marine worm; but this caused
alarm, and as the worm intended for food commenced to
burrow into, and I feared would destroy the new piece of
tube, I at once removed it. Panthals had, the while,
retreated into its burrow, but in course of about ten
minutes returned; forced its head slowly and carefully
through the mouth of the new tube, and proceeded to

178 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

repair the damage, lying on its back, and thus affording
me a splendid view of its operation through the open top
of the tank. Thus seen, the mouth of the new tube had
the appearance of a minute crater about + an inch external
diameter with walls barely #3,” thick at the edge. With
its head very slightly projecting through the opening, and
the two long palps bent over the edge ventrally, Panthalis
began moving the second pair of parapodia in a manner
quite different from the ordinary rowing or walking action
of such limbs; the motion, instead of being longitudinally,
was transversely to the body, 7.e., the extremities of the
right and left pairs of parapodia were brought together so
as almost to meet in the central line of the underside of
the body.

At intervals of two, or three seconds, the animal, for
several minutes at a time, continued thus to bring together
and to separate these limbs; the tips of the buccal tenta-
cles or bristles, too, at short intervals were brought together
and applied first to one point and then to another of the
tube, as though grasping and removing something from
place to place; and although no threads could with

certainty be seen (though I thought I saw the haziness of

a cobweb) I had not the slightest doubt that Panthalis
was thus weaving its tube; combing and weaving the
threads by means of the setze contained in these parapodia,
and arranging them with the buccal tentacles, an opinion
which I have since confirmed with another specimen,
when I saw the threads actually carried, as described
above. The process was repeated at different levels in
the tube which would account for the layers originally

noticed in the tube structures. Now and again the elytra |

took part in the formation of the tube, and by rising and
falling were used to expand its mouth, whilst occasionally
the edge was drawn in upon the animal and then forced

i
tr
\

TUBE OF PANTHALIS. 719

out again, possibly either with the object of shaking the
fine grains of mud amongst the threads, or of causing the
latter to become entangled. Rather long rests were made
between each spell of weaving, possibly to afford time for
further secretion of the threads. The action of the
“weaving feet,’ as described above, would result in an
arrangement of the threads approximately parallel with
each other, in a direction transverse to that of the length
of the tube, and upon examining, with the microscope,
part of the cobweb formed against the glass in the burrow,
it was found that this was actually the case. I ought,
however, to add that in subsequent instances this regu-
larity of direction was not always observable; the cobweb
being of a dense description similar to that formed by
some spiders.

The worm continued its weaving operations at intervals
during the morning until about noon, and then retired
into its burrow. It being impossible for me to remain
longer at Port Erin, I decided to convey Panthalis in its
tank to Sheffield, and there continue my observations.
This, with some difficulty was accomplished, and I was
fortunately able to keep the animal alive until the 17th of
September, a period of 23 days from the date of its capture.

Shortly after arrival in Sheffield the worm settled down
in its new quarters, and after clearing the openings of its
tube seemed habitually to lie with its head at one or other
of these points, the two long palps extended, or resting on
the mud externally. In this position it 1s most probably
watching for prey. On one occasion, when the palps had
remained thus outstretched for many hours, I feared the
animal was dead, and, in order to test it, gently raised one
of them with a quill, whereupon Panthalis attacked the
quill savagely with its Proboscis. —

During its life at Sheffield Panthalis afforded me an

180 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

opportunity of watching its method of mingling the fine
mud with the cobwebs, one of the points needful to com-
plete the process. This was done by the same limbs as
those used in weaving, the mud being swept into the
mouth of the tube by striking backwards, the edge of the
tube was also at the same time pulled in upon the animal
and forced out again as previously described. The dorsal
cirri of the 3rd pair of parapodia also appear occasionally
to assist by drawing fine mud into the tube. . It being
evident on the 17th September, that Panthalis was ailing,
I decided to remove it from its tube and examine it under
the microscope while still alive, as I had yet to ascertain
the source of the mucus like threads, and how they
were conveyed to the ‘‘ weaving feet,” Thanks to Miss
Buchanan’s admirable paper on Hupolyodontes cormishi in
the Quart. Journal of Microscopical Science of January,
1894, in which she briefly describes the members of the
sub-family Acoétide, and gives the bibliography on the
subject, I was already aware of the existence of Hisig’s
‘‘spinning glands’’ in the parapodia of Polyodontes, and
consequently prepared to expect something similar in
Panthalis.

-This particular living specimen afforded no evidence on
that point, but by observation of others, and also by
dissection of dead specimens, I.am convinced that these
glands, which exist in all the parapodia of Panthals,
except the anterior eight pairs, are the source of the
mucus-like threads. The glands correspond very closely
with those of Polyodontes maxillosus described and figured
by Hisig (Monographie der Capitelliden), and their laige,
bronze-like coils, seen on opening the body of the animal
either from above, or by transverse section, are most
striking objects. In the latter view the chetal sac when
opened, reminds one of a lock of hair, the long, exceed-

TUBE OF PANTHALIS. 181

ingly fine capillary chet, lying compactly side by side,
readily separable with a dissecting needle. The extensions
from these coils can easily be traced to the cleft between
the upper and lower lobes of the parapodia, from which
points I have, more than once, seen the white, tube- -
forming threads to issue. Whether these threads are
themselves prolongations of Cheete, or are fabricated by
sland secretions, forced through the chete, which are
shewn as tubes by Hisig, is a point to which I hope to
give further attention.

It may be noted that the dorsal lobes of the parapodia
endowed with spinning-glands, are much wider than those
of the preceding limbs.

On the remaining point, viz., how the threads are
passed forward to the ‘‘ weaving feet,’ I fortunately gained
a clue from this living specimen. Kinberg in his des-
cription of Panthalis oerstedi (‘‘ Fregatten Hugenies
Resa”’ Zool. Annulater) specifies the sete as of three
kinds ‘‘ subulato-serrulate, bipennato-penicillate, and
aristate,’ and states that the first-named occupy the
lowest position as regards the parapodium ; the aristate
sete, the medium, and the bipennato-penicillate the
superior or dorsal; the action of the medium and superior
setze 1s described as perpendicular, and that of the inferior
sete, as horizontal. These sete, as well as the animal
itself, are figured in the work above referred to, but those
observed in my specimens appear to be a combination of
the setze given both for Panthalis and Hupompe grubet,
and correspond most closely with the latter. (Possibly
Kinberg got his figures mixed.) None of these figures,
however, give quite the correct impression of the “ bipen-
nato-penicillate’”’ setae of Kinberg. When in action, it is
really that of a minute brush, in which the bristles have
flexible attachments round the tip of a long central shaft,

182 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

somewhat like the ribs of an umbrella which has been
turned inside out. Often these ribs are partially closed
up, and shew no movement, but I have several times seen
them when at work opening and closing, probably by the
pull of the threads and the resistance of the water. It
should be noticed that the ‘‘ brush-like ” sete are attached
only to the parapodia possessing “‘ spinning-glands”’ and
their use is at once obvious. All the sete in Panthalis
proceed from the ventral lobes of the parapodia; the
position of these brush-like bristles, as described by Kin-
berg is uppermost; they are capable of very considerable
extension and also of being retracted almost entirely
within the body of the animal; the threads from the
‘spinning gland”’ issue from the cleft between the upper
and lower lobes of the parapodia, 2.¢e., just against these
sete, and they are caught-up and carried outwards and
forward by the action of these brushes assisted, to some
extent, in the latter movement by other sete. I have
seen this actually take place, and on examination of dead
specimens the observation is confirmed, the brush-like
sete being frequently found directed forward with masses
of threads entangled in them. ‘

The parapodia possessing “spinning glands” are en-
dowed with all three kinds of setee mentioned by Kinberg ;
but the six pairs between the last ‘‘ spinning gland ” and.
the ‘ weaving feet’”’ have two kinds only, the brush-like
setee being replaced by a second set of serrulate sete
similar to those figured by Kinberg for Hwpompe.

The foregoing, I think, describes fully the formation of a
single thin tube; but it will be remembered that the
tube inhabited by Panthalis 1s quarter of an inch thick,
composed of many apparently parallel layers, formed one
inside the other, and I therefore still had some difficulty
in understanding how this strong, heavy structure could

TUBE OF PANTHALIS. 183,

be produced. An examination of the hardened and
embedded sections of tubes first made; and of fresh
sections of tubes in their natural condition; and an exter-
nal examination of the tubes themselves under running
water (bearing in mind the bursting action possessed
and exercised by the forepart of the animal’s body) afford,
I think, a simple explanation.

The sections shew that the layers, although apparently
parallel, do not run horizontally the full length of the
tube, but all curve outwards, and it is due to this fact,
that Panthalis is able to construct a tube of so great
thickness. The process, most probably, is as follows :—
A moderately thin tube is-first formed, consisting of cob-
websandmud. In consequence of the layers being added
internally, the tube would soon become inconveniently
small, and the animal thereupon burst the anterior portion
(as | saw it burst its original burrow) by expansion of the
forepart of its body, thus throwing the mouth of the tube
backwards. It then proceeds alternately with the addition
of more internal layers and the bursting process, by which
means the free ends of the internal tubes are successively
thrown outward and the layers are made to take the
outward curve above referred to, the final effect being that
of a series of hollow truncated cones, one inside another.

That this is the case is proved both by the fresh section
in which the layers can be separated, and by an external
examination of the tube masses under running water. By
the latter means the different tubes can be partially folded
back, but as the split side is always made good in the
formation of the new inner tube to which it is attached,
these tubes can only be folded back for a portion of the
circumference. The long straggling mucus threads which
always accompany the tubes are probably formed by the
animal when exploring its surroundings.

184 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

In this paper I do not profess to have gone minutely
into details of the anatomy of Panthalis, but in the course ~
of my observations, one or two points of interest have come
before me, and a brief reference to them may be useful.

The sight of the animal is evidently very good, and a
remark as to the structure of the organs of sight will be
interesting. The eyestalks, as also the cephalic mass to
which they are attached, are brownish-red in colour, and
have at their tips a rounded papillated appearance. . Hach
eyestalk is faced with a clear lens, the curve of which is
occasionally somewhat acute, though this seems to vary,
and possibly the animal may have some means of adapting
it to circumstances.

The structure and action of the Proboscis also deserve
a passing note. Asis usual in annelids, this organ 1s very
often protruded in the act of dying, and thus affords an
opportunity for observation of its action. In Kinberg’s
figures of Panthalis and Huponwpe the proboscis is shown
protruded, but the peculiar structure of this organ is
hardly noticeable. My specimens agree more closely with
his figure of Hupomwpe than with that of Panthalis. As
~ described by Claparéde (Les Annélides Chétopodes du Golfe
de Naples, 1868) the proboscis is divided into two parts,
a longer inferior portion consisting of a cylinder of
somewhat ordinary form, and a short superior portion,
which carries the jaws and is fringed with papille, the
central ones of which, both dorsally and ventrally, are
considerably elongated. The dorsal one being about = of
an inch long (in my large specimen) forms a tentacle.
The action of the organ is a double one, and as follows :—
the lower part only of the proboscis is first protruded,
either for the purpose of burrowing in the mud (as I have ~
seen it used) or in search of food. In the latter case, the
central dorsal tentacle attached to the margin of the

TUBE OF PANTHALIS. 185

proboscis simultaneously appears resting in a fold on the
upper surface of this lower part. This tentacle doubtless
ascertains whether food is within striking distance in which
event the second portion of the proboscis is brought into
action, seizing its prey by means of the four savage-looking
fangs, assisted by the smaller teeth with which the animal
is endowed.

The campanulate structure of the elytra is interesting,
and raises questions in my mind as to how far these are
organs of respiration specially adapted to expose as large
a surface as possible to the action of the water, in view of
the confinement in which Panthalis usually lives.

From specimens which I have had under observation
during several months I should gather that, although
Panthalis does occasionally desert its tube and make new
ones, as suggested by Prof. Herdman, (see British Asso-
ciation Report, Oxford, 1894) this is not a very frequent
occurrence, and that when once the worm is well-settled its
general habit is to rest with its head at the mouth of its
tube watching for its victims, an opinion apparently
supported by the facts that two tubes dredged in September
last, have each one end hard and leathery, as though it
had been long buried and hardened by absorption of
excreta or other matter, and that the complete formation
of such tubes seems to be a work of time. The animal
appears always to leave its tube when about to die, which
will account for many empty ones being brought up in
the dredge.

As to its mode of locomotion, Panthalis usually walks
or creeps upon the mud. I have only once seen it swim,
which it did in a clumsy way, by moving its elytra and
body on an occasion when the water had become foul and
unbearable.

Throughout this paper, I have, for the sake of con-

186 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

venience, referred to the threads secreted by Panthalis as.

‘““mucus-like”’ but I don’t intend thereby to express an
opinion as to the nature of the material of which they are
composed. ‘They are secreted by Hisig’s spinning elands
and appear to be very similar to those formed by Polyodontes
maxtllosus, which Hisig describes as ‘‘ chitinous.”” They

are very slightly, if at all, soluble in boiling caustic potash.

_ As explained at the outset, my chief object in these
observations was to ascertain whether Panthalis. made
the tube which it inhabits and if so how? and I think I
have satisfactorily answered both questions.

The tubes of the Acoétide, from the absence of any
membranous lining, and their general construction, would
appear to be unique amongst annelids. In this connexion
see Quatrefages’ reference to Acoétes pleet, (Histoire
Naturelle des Annélés, I, p. 216), Hisig’s account of Polyo-
dontes mazxillosus, and Hhler’s description of Huarche
tubifex (Florida Anneliden, pp. 54-6). No minute
description has previously been given, but from the
similarity of their organs, it seems probable that the tubes
of all these worms will more or less closely resemble
each other.

In closing this paper I would take the opportunity of
expressing my great appreciation of, and gratitude for the
advantages afforded me at the Port Erin Biological Station,
to which the success of my experiments has been largely
due. Absolute stillness was a most important factor, and
Iam greatly indebted to Prof. Herdman, both for the steps
which he took to ensure this, and also for very much
valuable advice and assistance. To him this paper is
really due, as excepting for his kindness in making special

journeys with me on August 25th, and September 30th, |

1894, to the ground where Panthalis is to be found, these
observations would have been impossible.

io, 1:

ie, 2.

ETO iia).

TUBE OF PANTHALIS. 187

EXPLANATION OF PLATES.

PuaTe IX.

Longitudinal section of wall of tube of Panthalis.
Sketch (shghtly enlarged) made from portion of
tube sectionised under water to shew the series of
layers thrown outwards. The darkest part (0) is
the inside of the tube.

Sketch of Panthalis in its burrow, shewing it in
a very frequent position, viz., twisted so as to
expose the front ventral, and posterior dorsal parts.
The palps extend in front; the buccal tentacles
almost ineet in the form of a V, and the ventral
bloodvessel, with its broad front, runs horizontally
towards the rear. The cobweb first formed, 1s
indicated to the right of the burrow, and at the
opposite end is seen the remnant of the cobweb,
by means of which (mingled with mud) the new
tube was made. The opening of the new tube is
shewn as a little mound (a) at the left of the
picture. The old tube forms the mass above the
burrow.

Enlarged sketch. Ventral view of anterior por-

tion of Panthalis shewing the two long palps (a)

(ends broken off); cirri of first pair of parapodia
(b); eyes and eye-stalks (c); between which is
seen the tip of the median small tentacle (d), and
in front the small paired tentacles. The ‘‘ weaving-
feet’ (e), with bristles, and ventral cirri or buccal
tentacles, directed across the mouth, followed by
other parapodia. The ventral bloodvessel (/)
with its broad square termination.

188 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

ie. de

Fig. 2.

Fig. 3.

Fig. 4,

Fig. 5.

PLATE X.

Sketch from micro-photograph of a ‘‘ spinning- —
foot”’ shewing all three kinds of sets, and two
‘““mucus”’ threads (m) entangled in the brushes,

Sketch from micro-photograph of a ‘“‘ weaving-
foot’? shewing its peculiar, very finely serrated
sete. v. Ventral Lobe. d. Dorsal Lobe.

Diagram. Ventral view of lateral portion of
Panthalis, shewing arrangement of sete in a
‘“‘ spinning-foot.” To the left are shewn all three
kinds. The ventral and dorsal sete have been
removed from the second foot; the dorsal ‘‘ brush-
like’ setee only are shewn in the third foot; and
‘“ spinning-gland ”’
(e) with threads issuing and carried away by the -
‘“brush-like”’ setae (d), are seen in the 4th.

Sketch from micro-photograph of ‘“ spinning-
gland” dissected out and partly uncoiled. Shewing
the strong ventral (a), and weaker dorsal (0)
acicules ; also the stout median sete (c), and the
delicate brush-like sete (d). The latter will be
seen to emerge close to the mouth of the “ spin-
ning-gland” (e), which has, however, got pulled
slightly out of position in dissection.

Sketch from micro-photograph of front part of
above more highly magnified. In it the “‘ brush-
like’’ setee (d) can be distinctly traced passing by
the mouth of the ‘‘ spinning-gland ”’ (e).

Proboscis (lateral view) shewing the palps, which
are usually straight, but have curled up in dying.
The two portions, viz., the trunk (a) of Proboscis, _
and the jaws (6), with long median tentacles (c),
the large fangs (d) and smaller teeth. (Much
enlarged.)

the position and form of the

189

OBSERVATIONS upon the POLLEN TUBE.
By A. J. Hwarrt, B.Se.

1851 EXHIBITION SCHOLAR. LATE DEMONSTRATOR OF BOTANY IN
UNIVERSITY COLLEGE, LIVERPOOL.

With Plate XI.
[Read 14th December, 1894.]

THE following paper gives the results of an investigation
on certain points in connection with the pollen tube,
which was suggested to me by Professor Reynolds Green,
F.R.S. Whilst investigating the zymogenic activity of
the developing pollen tube Professor Green came to the
conclusion that the enzyme was excreted from the apex
of the tube in the form of solid particles of zymogen,
and certain appearances seen in the pollen tubes of
Narcissus led him to infer the presence of a definite and
permanent excreting pore in the apex of the pollen tube.
At his suggestion I attempted to determine the nature of
this pore and whether its presence was a constant and
normal feature of the pollen tube.

On examining the apices of stained and mounted pollen
tubes, under high powers (;'s immersion) a few on each
slide were seen to shew very minute terminal oval or
rounded pores and in some cases a small amount of gran-
ular material which had evidently been extruded by the
tube through the pore could be seen still adhering to the
outer surface of the apex of the pollen tube. These pores
could be satisfactorily distinguished only in normal pollen
tubes of Narcissus, Tulipa, Lilwwm and Scilla though the
apices of the pollen tubes of several other plants, Lathyrus,
Brassica, Cheiranthus, Vacconiwm, Mercurialis, etc., were
similarly examined. ‘The question which needs to be
answered is whether this pore is permanent and the path

ee

190. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

by which the zymogen particles are extruded, or whether
it 1s a temporary opening made by the particles as they
escape, which, being of recent formation, has not yet
closed up and disappeared.

The fact that it is only in relatively few cases that the
pore can be seen would seem to indicate that the latter 1s
the more probable view. Supposing, however, that the
pore is a permanent structure opening only under the
action of an internal pressure caused by a local or general
increase of turgidity, the lips of the pore coming so closely
together that the latter is obliterated and becomes invisible
after turgidity is relieved, then it would only be in a few
cases that we should see the pore open and hence visible
in a stained and mounted preparation. .

In order to determine whether this latter view was correct —
or not experiments were performed on pollen tubes by
varying their turgidity and then examining them. Pollen
tubes of Narcissus which had germinated in a sugar
solution of normal strength were placed some in a strong
sugar solution and some in a very dilute solution; strong
and very dilute solution of potassium nitrate were also
used. An hour after, a quantity of a saturated watery
solution of picric acid was poured over the pollen tubes
in order to kill and fix them in the particular condition in
which they might then happen to be. The pollen tubes
placed in the dilute solution have their turgidity increased,
those in the concentrated solution have their turgidity
diminished, hence in the former case if the primary sup-
position be true the pores should all be open and visible
and in the latter case they should be all closed and
invisible. Microscopical examination, however, shews _
that the pores are no more patent in the former case than
in the latter. It was found, however, that after the
pollen tubes had been kept for some hours under conditions

OBSERVATIONS UPON THE POLLEN TUBE. 191

involving increased turgidity the pores are somewhat more
numerous and readily visible.

On examining living pollen tubes of Narcissus under
the microscope the actual extrusion of solid granules
through the apex of the tube may be seen. This normally
is a very slow and gradual process the granules appearing
to work through the wall of the tube and not to pass
through a permanent aperture (PI. XI. fig. 10). Only a
few particles escape at a time and these form a little clump
of finely granular material which remains for a short time
adhering to the outer surface of the pollen tube. After
the particles have escaped the pore closes up after them
and disappears, the walls of the opening fusing together
(fig. 11). Changes of turgidity of moderate intensity
produced by irrigating the living tubes with concentrated
and dilute sugary solutions exercise no immediate effect
upon the formation of the pores.

In some cases a very large and wide terminal opening
is formed, which may be from 1} to 2 of the diameter of
the pollen tube (fig. 12). Through this opening a large
part of the contents of the pollen tube escape whilst the
water which can now readily enter the pollen tube through
the wide terminal opening rapidly disorganizes its con-
tents, the granular debris thus formed shewing marked
Brownian movement.

The opening thus formed differs from a pore only in its
much greater size and in its being unable to close when
once formed. ‘The same causes which produce the pore
also produce by a more violent action these apical ruptures
of the pollen tube. The internal turgidity of the pollen
tube presses the limiting layer of ectoplasm tightly against
the cell wall and the latter stretches until its elastic
tension is equalled by the internal hydrostatic pressure.
The apex of the pollen tube being softer, more ductile and

192 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

less elastic than the side walls of the tube is stretched by
this internal pressure and the stretching being rendered
permanent by growth, the pollen tube grows in length.
It may happen that the soft ductile region is not terminal
but remains somewhat lateral in position. In such cases
the pollen tube grows spirally and the closeness of the
spiral is determined by the amount of deviation of the
softened apical region from the apex of the tube. If the
terminal softer apical region divides into two more laterally
placed areas the pollen tube will fork at this spot. The
branching of the pollen tube, which is of rare occurrence
and which apparently takes place only under abnormal
conditions, seems, except in certain cases to be mentioned
later, to be always of apical origin and is hence dicho-
tomous though it may happen that one of the branches
becomes larger than the other and continues the growth
and direction of the unbranched tube.

In certain cases the growth of the tube having been
checked owing to a diminution of turgidity the wall of the
terminal apical portion being no longer continually
stretched becomes thicker, shews faint lamellation and is
evidently softer and contains more water than the older
part of the tube. It sometimes happens that not only
the wall of the extreme apex but also of the terminal
portion of the pollen tube remains soft and ductile, and in
such cases any increase in the internal turgidity causes a
swelling or bulb like dilatation to be formed at the apex
of the tube. These apical dilatations are found most
abundantly in old cultures, though they may be formed
as soon as the first protrusion of the endosporium takes
place. The formation of these swellings seems to be the -
final stage in the life history of the pollen tube for in no
case has a commencement of the growth and elongation
of the tube been seen after they have once been formed,

EE Ee ee ee ee ee
- = * ’ — ‘ “TA - ig Ss Tr

OBSERVATIONS UPON THE POLLEN TUBE. 198

nor does any normal extrusion of granules appear to take
place through such swollen apices.

When the pollen grain germinates upon the stigma this
stage will occur when the apex of the pollen tube reaches
the surface of the embryosac and the apical softening
which then takes place will readily allow of the extrusion
and escape of the male pro-nucleus from the enlarged and
swollen apex of the pollen tube.

To sum up, the reasons for concluding that the minute
pore, which may be seen in the apex of certain pollen
tubes, is not a permanent structure, but opens on increase
and closes on decrease of turegidity, are:

(1) The pore may be to one side of the apex instead of
being terminal, 7.e.,1t does not always occupy the
same position.

(2) More than one pore, as many as two or three, may
be present in the apex of one tube.

(3) After the granules have escaped, the pore totally
closes and disappears and may next time be formed
in a different situation.

(4) The bulbous enlargements formed on the ends of
the tubes, which are swellings of the softened apex
could not be formed if a permanent pore opening on
an increase of turgidity taking place were present
in the apex of the pollen tube.

It is worthy of notice as instancing the extreme care
needed in interpreting the appearances seen In an investi-
gation of this kind that once or twice an appearance was
seen in the apex of a pollen tube exactly resembling an oval
opening or slit with regular and well marked lips. This
was really a depression or crease in the apex of the tube,
the margins of the depression looking exactly like the lips
of an oval slit. Closer examination shews the endos-
porium to be continuous between the margins of the

194 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

depression and simple pressure on the cover glass may
cause the filling up and disappearance of the crease.

In performing prolonged culture experiments with
pollen tubes it 1s found that the culture medium soon
becomes filled with Bacteria, fungal hyphe and Yeast
cells and that the development of these organisms injur-
iously affects the development and normal growth of the
pollen tubes. During the course of the experiments
various attempts were made to obtain pure cultures but
without success. The effect of adding to the sugary
medium traces of mineral acids, of salicylic acid, etc., was
tried. In all cases 1t was found that a percentage of acid
sufficient to almost entirely inhibit the development of
Bacteria was insufficient to prevent the formation of
fungus mycelia or the appearance of Yeast cells and yet
prevented the germination of the pollen grains. It is
interesting in this connection to notice that cultures of
pollen soon became acid and hence in cultures where a
vigorous and rapid growth af the pollen tubes takes place
the excessive development of Bacteria is to a certain
extent checked. Similarly using other poisonous sub-
stances (mercuric chloride, sodium arsenate, alcohol, etc.)
it was found that any reagent powerful enough to inhibit
the development of Yeast, Bacteria and Fungi was also
sufficiently powerful to inhibit the germination of the
pollen grain and the formation of the pollen tube.

By adopting certain precautions a fairly pure culture of
pollen tubes may be obtained. The sugar solution should
be boiled before using, all vessels and instruments used
should be previously heated to 150°C and the pollen

should be taken not from dehisced anthers but from_
anthers which are just about to open, whilst the tubes

containing the culture should be plugged with sterilized
cotton wool. In this way cultures may be obtained in

OBSERVATIONS UPON THE POLLEN TUBE. 195

which the disturbing influence due to the presence of
foreign micro-organisms may be considerably reduced.
Fungal hyphe and Yeast cells being absent whilst
Bacteria are for a time at least relatively few in number.
It is impossible to obtain any fairly large quantity of
pollen which shall at the same time be entirely free from
the spores of Bacteria and any attempt to destroy these
would necessarily involve the destruction of the pollen
grains also. ‘The spores thus introduced though perhaps
at first but few in number soon by their rapid development
and multiplication spoil the cultures, but for the first two
or three days and especially if the tubes are exposed to
bright sunlight the numbers present are fairly low.

Difficult as it is to obtain pure tube cultures of pollen
and to keep such free from the disturbing influence of
Bacteria, etc., yet this is the only way by which a pro-
longed experimental culture can be made; a sample of
the developing tubes being examined from time to time.
Here of course chances of error owing to the necessary
manipulation are introduced and the sample may not,
unless two or three are taken, accurately represent the
condition of the bulk of the culture.

Unringed slide cultures on the other hand rapidly
evaporate and if ringed with vaseline to prevent evapora-
tion, the supply of oxygen which the enclosed fluid
contains is soon used up and the growth of the pollen
tubes ceases. The rapid stoppage of growth which thus
occurs is due to the cessation or diminution of the tur-
eidity of the pollen tube, probably owing to a diminished
percentage of osmotically active substances, organic acids,
etc., in the cell sap and this again is due to the deficiency
of oxygen.

When attempting to inhibit the development of Bacteria
by exposing tube cultures of Narcissus pollen to the action

196 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of direct sunlight, it was noticed that at the same time
the rapidity of growth of the pollen tube was also checked.
To determine this retarding influence of direct sunlight
upon the growth of the pollen tubes more accurately
detailed comparative experiments were made. A culture
solution containing fresh Narcissus pollen was divided
into two equal portions, one of which was placed in a tube
covered externally by several layers of black asphalt
varnish and the other in a similar, but clean, glass tube.
On taking the temperature of these tubes when exposed
to sunlight it was found that the temperature of the tube
covered with asphalt was higher than that of the clean
glass tube but on pasting thin white paper on the more
absorptive black surface the temperature of the two tubes
became approximately equal. Thus in tubes half filled
with sugar solution :
1. Asphalt tube exposed to sunlight, temperature was
ESI Oe
2. Clean glass tube exposed to sunlight, temperature
was 20°5°C.
3. Asphalt tube covered with white paper exposed to
sunlight, temperature was 20°6°C.
4. Asphalt tube covered with white paper in ae
temperature was 14°5°C.
5. Clean glass tube in shade, temperature was 14°8°C.
In (8) when exposed to sunlight the temperature is only
‘1 higher than in (2) and in the shade or at night the
temperature in the darkened tube is ‘2 or ‘3 lower.
These differences of temperature are too slight to be able
to markedly affect the growth and elongation of the pollen
tubes. :
A much more difficult matter was to obtain suficiontie
long periods of exposure to sunlight. Time after time an
experiment started in bright sunlight was spoilt by the

OBSERVATIONS UPON THE POLLEN TUBE. 197

sky becoming overcast with cloulds. It was only after
dozens of experiments had been made during the summer
months that satisfactory results were obtained. The
experiments were performed with pollen of Narcissus,
Chetranthus and Lathyrus. In each case after a certain
length of time samples from each culture were taken and
the number of pollen grains, and the number and average
length of the pollen tubes in each estimated and compared.
_ The following are illustrative examples of the results
obtained :

Narcissus examined after 14 hours exposure.

(1) In darkness. Average length of tubes 4 to 6 times

diameter of pollen grain.

(2) In diffuse daylight. Average length of tubes 6 to

10 times diameter of pollen grain.

(3) In bright sunlight. No tubes formed.

In a few cases in (3) the commencing protrusion of the
pollen tube was seen but not more than this. HWxamined
at the end of the second day (t.e., after exposure for two
days and one night) a number of tubes were seen to have
been formed. These averaged only half the length of the
pollen tubes of (2) and compared with (2) 40 per cent. less
tubes were formed to the same number of pollen grains.
The major part of this growth and germination takes
place during the period of darkness necessarily interposed
between the two periods of exposure to sunlight. The
same experiments with pollen of the wallflower gave
similar results. Thus exposed to sunlight but few
germinated whilst in diffuse daylight and in darkness
a larger percentage germinated, the average lengths
of the pollen tubes being respectively from 2 to 4, from
8 to 10, and from 7 to 9 times the diameter of the pollen
erains when examined after being in the culture solution
for a night anda day. Pollen of Lathyrus odoratus after
10 hours, during 5 of which the sun was shining :—

198 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

(2) Direct sunlight. Length of tubes 8 to 4 times
diameter of pollen grain. About 10 p.c. of pollen
grains germinated.

(0) Shaded from sualight. Length of tubes 6 to 8 times
diameter of pollen grain. 15 p.c. germinated.

(c) In darkness. Length of tubes 6 to 8 times diameter

of pollen grain. 20 p.c. germinated.

After 36 hours, 12 hours full sunlight, 14 hours daylight
but no sun, 10 hours darkness and twilight.

(a) Direct sunlight. Average length of tubes 7 to 10

times diameter of grain.

(0) Shaded from light. Average length of tubes 15 to

18 times diameter of grain.

(c) In darkness. Average length of tubes 14 to 16

times diameter of grain.

In these tubes exposed to diffuse daylight the tempera-
ture is distinctly lower than in those exposed to the sun,
and hence the growth of the pollen tubes, in such cultures,
is not so rapid as it would otherwise be; but on the
other hand the excessive development of Bacteria, etc.,
checking the growth of the pollen tubes is less marked in
the cultures exposed to diffuse daylight than it is in the
darkened tubes. These two antagonistic influences partly
neutralize each other but the influence of the lower
temperature in retarding the growth of the tubes is much
more marked and produces a direct and immediate effect.
Hence in (0) the growth of the tubes would be greater
than in (c) were the temperatures equal in the two cases.

These results show that direct sunlight retards the
growth of the pollen tube and may inhibit the germination
of the pollen grain whilst exposed to weak diffuse day-
light. The rapidity with which the pollen tube elongates
is slightly greater than it is in absolute darkness. In

nature, a pollen tube is neither exposed to direct sunlight ~

OBSERVATIONS UPON THE POLLEN TUBE. ~ 199

nor is it in absolute darkness, the stigma being as a
general rule shaded from the sun whilst through the
partially transparent tissue of the style into which the
pollen tube burrows a certain amount of light can
penetrate.

In aerial plants light of any intensity retards growth
because it promotes transpiration and hence lessens that
turgidity of the growing organs and regions which is so
necessary a factor in the continuance of growth. In the
case of a pollen tube, however, which is growing immersed
in a nutrient solution it is evident that hght can exercise
no such influence. The effect of direct sunlight in
retarding growth is due to a direct action upon the
protoplasm of the pollen tube, causing it to assume a
condition of ‘immotility ” and hence causing a cessation
of tureidity and stoppage of growth. Darkness exercises
a similar but very much weaker influence. The motility
of the protoplasm of the pollen tube and the turegidity of
the latter are most pronounced when exposed to weak
diffuse daylight. The four factors which influence the
rapidity of growth of the pollen tube are; the nature and
strength of the nutrient medium, the relative amount of
oxygen present, the intensity of the light to which the
culture is exposed and the temperature at which it is kept,
the optimum being between 20° and 30°C.

When examining two and three days old cultures of
Narcissus pollen it was noticed that in some cases the
pollen tubes had become septate. The septa were in the
form of rather thick transparent transverse partitions and |
were few in number, there being rarely more than four or
five in one pollen tube. At first sight the appearance of
these septa seems to indicate that the pollen tube had in
such cases undergone cellular division and become a
multicellular filament, for the septa may not only divide

200 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the tube into separate segments but may also segment

its protoplasmic contents. They may be formed at the —

base of the tube where the protcplasm has all flowed away
towards the apical part of the tube, as well as in regions
still filled with protoplasm.

The formation of these septa seems to be due to a
pathological change in certain regions of the wall of the
pollen tube. They are shewn best after staining with
picric-aniline. If left staining in this reagent for a
limited period, the walls and septa are stained blue whilst
the protoplasmic contents owing to the picric acid
penetrating first are stained yellow, though if left in the
stain for a longer time they become yellowish blue. By
this method of staining the ingrowths are rendered
very conspicuous. The septa are seen on testing micro-
scopically to be composed of cellulose. Both the walls
of the pollen tubes and the septa formed from them are
too delicate to allow the sulphuric acid and iodine test
to be employed, the acid either shrivelling up the tubes
or totally destroying them. On treating with chloro-zine
iodine solution the walls of the tubes swell up and turn
blue, the cavity of the tube being almost obliterated and
the septa disappearing. If, however, a septum is watched
whilst this reaction is taking place the tube is seen to be
at that point solid and blue throughout. The refractive
indices of the walls and septa are after treatment with
chloro-zince iodine solution similar though formerly they
were not so. The septa being portions of the cell which
have already swollen do not swell any further whereas the
cell wall under the action of the reagent does swell and
its refractive index is now the same as that of the septum.
Cupr.-ammonia dissolves both walls and septa. ;

On examining a series of living pollen tubes the origin
of these septa can readily be made out. ‘They arise in the

OBSERVATIONS UPON THE POLLEN TUBE. 201

form of small ingrowths from the wall of the tube. A
pair of these may be formed opposite each other and |
erowing together form a complete partition but as a
general rule the septum is formed as an ingrowth from
one side of the tube only. The ingrowths swell up,
commonly becoming tuberculated and may form irregular
amorphous masses blocking up the cavity of the pollen
tube. They are formed by the localised imbibition of
water causing localized swellings and the fact that these
are always found internally and never externally shews
that the inner part of the wall of the pollen tube is softer
less resistant and more capable of excessive imbibition
and of undergoing a partial mucilaginous modification
than is the outer layer.

Exactly similar partitions though neither so numerous
nor distinct as in Narcissus, were found in normal cultures,
of one or more day’s growth, of all the pollen grains
examined, viz., Cheiranthus, Vaccinium, Tulipa, Lathyrus.
They are therefore of very general occurrence and though
they are to be regarded as pathological growths are never-
theless in same cases apparently of importance. Thus in
an old tube of some length in which the basal part and
the cavity of the pollen grain itself are alike devoid of
protoplasmic contents it is evidently an advantage that
this portion should be, as it often happens in Narcissus,
eut off from the living apical portion of the tube and hence
the maintenance of turgidity necessary for the elongation
and growth of the pollen tube restricted to that portion
_ of the tube in which the cell-wall is still bounded internally
by a lining film of protoplasm. On the other hand the
formation of a septum across the middle of a tube cutting
off a portion of the protoplasmic contents cannot but be
disadvantageous and this is especially the case when, as
sometimes though rarely happens the part of the proto-

Fs 202 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

plasmic contents thus cut off contains the generative
nucleus or nuclei. The formation of these ingrowths is
frequently preceded by a general thickening of the wall of
the pollen tube; and is most marked in Vacconewm where -
the wall may more than double its previous thickness.
The conditions most favourable for the production of
the septa were investigated with pollen of Narcissus by
cultivating 1t in sugary solutions of different strengths.
In very dilute solutions the grains become very turgid
swelling up considerably. In some cases the limit of
stretching being reached the coats burst and the contents
escape and become rapidly disorganised but in other cases
only the exosporium breaks and the entire endosporium
escapes, enlarges considerably and assumes various irregu-
lar shapes. Since this enlargement is beyond the limit ~
of stretching growth must have taken place over the
entire surface of the endosporium, the irregular shape
which the latter assumes being the result of attempts to
form a pollen tube, the wall where a protuberance is
formed being softer and more ductile than elsewhere.
i No proper pollen tubes are formed. In a strong sugary
solution the grains swell but little and become very
transparent. ‘The protoplasm is in most cases shrinks
more or less from the wall of the pollen grain and owing
to the diminished turgidity of the latter the number and
the rapidity of formation of the pollen tubes is much
diminished. The shrinking of the protoplasmic contents
of the tubes may be so marked, that some of the starch
grains are squeezed out of the shrunken protoplasmic |
mass and lie in the space between it and the wall of the
tube. This is especially the case when tubes grown in
dilute solutions are placed in concentrated sugary solu-
tions. The branching of the pollen tubes, never very
abundant, occurs oftenest in strong sugar solutions, and

i

OBSERVATIONS UPON THE POLLEN TUBE. 203

seems always to be, or to become, more or less dicho-
tomous. It may be dichotomous from the commencement
but in other cases the branch commences as a swelling a
little below the apex, and is hence originally lateral,
afterwards lengthening so rapidly as to rival the length of
the original apical portion and hence becoming dichoto-
mous. It is rarely that a tube branches more than once
and I have never seen more than two branchings occur
in the same tube.

Pollen grains of Mercurialis may emit two or even three
separate tubes all arising close together but distinct to
their bases. In other cases three tubes are formed by the
pollen grain forming the primary tubes and one of these
branching dichotomously. ‘The formation of two tubes

from one grain is of fairly frequent occurrence but these

tubes are only very rarely seen. ‘The formation of cellu-
lose ingrowths is in the case of Narcissus most frequent
is sugary solutions of some strength whereas since the
ingrowths are really irregular internal swellings produced
by the excessive localized imbibition of water, we should
expect to find them more abundantly formed in weaker
than in strong sugar solutions. On the other hand the
percentage of solids in the fluid, contained in the pollen
tube and touching the inner side of the wall, will be less
as compared with that of the external medium when in
a strong, than when in a weak, sugary, solution. Hence
at any spot where the sugar penetrates in less relative
proportion to the water that its does elsewhere and where
the internal layers of the wall of pollen tube are soft and
readily capable of imbibition an internal swelling in the
form of an irregular cellulose ingrowth 1s produced.

It sometimes happens that wrinklings of the wall of the
tube simulate an appearance of septa. These form thin
lines running generally diagonally across the tubes and

204 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

altering in position as they are focussed. They are hence
readily distinguishable from true septa. |

The foregoing observations were made in the Botanical
Laboratory of University College, Liverpool.

1 | ; q
)
{
|

EXPLANATION OF PLATE XI.

Figs. 1—6, 10—19, Narcissus; 7—9, Mercurialis; 20,
Vaccommum ; 21, Cheiranthus.

Fig. 1. Commencement of dichotomous branching.

Big, 2, 5s ,, lateral =

Fig, 3. Later stage of 2.

Fig. 4. Young plasmolysed pollen grains, shewing extru-

sion of starch grains.
Fig. 5. Forms produced in dilute sugar solution.
Figs. 7—9. Formation of normal and branched pollen tube.
Fig. 10. Extrusion of zymogenic granules from apical pore.
Fig. 11. Apex of same tube three hours later.
Fig. 12. Apex of pollen tube which has just ruptured.
Fig. 13. Swollen lamellated apex of pollen tube which has -
ceased to elongate. i
Fig. 14. Apex of old pollen tube shewing globular aii
tion, formation of ingrowth and septum-like
crease.
Fig. 15. Same pollen tube under low power.
Fig. 16. Formation of septum along with turgidity of

basal portion of tube causing an attempt at
branching.
Figs. 17—18. Commencing ingrowths.
Fig. 19. Lamellation of wall and Ingrowths, [7; Im-
| mersion ] .
| Fig. 20. Formation of septa and thickening of wall.
Fig. 21, Septa enclosing reproductive nucleus.

205

NOTES on a COLLECTION of SPONGES from the
WEST COAST of PORTUGAL.

By R. Hanitscn, Ph.D.

With Plates XII. and XIII.
[Read May 17th, 1895. |]

THE following pages contain a description of a small, but
interesting collection of Portuguese Sponges, which were
sent to me by Professor Paulino de Oliveira, Coimbra.
They are all marine, with the exception of one, Huspon-
gilla lacustris, Autt., the only species of Fresh-water
Sponges thus far obtained from the Iberian Peninsula.

I have not, in all cases, succeeded in specific indentifi-
cation, and with some of the forms I have not even
attempted it. Many of the specimens which I received
were mere fragments cut from larger specimens, and this
often made identification more difficult, especially with
the Horny Sponges. In the case of the genera Halichon-
dria, Remera and Chalina, I did not seriously attempt
specific identification, as I consider a vast number of
species of those genera to be quite insufficiently defined,
and before somebody takes the trouble to work out those
genera, I do not think it of much use to attempt specific
identification.

The collection comprises twenty-eight forms. Two of
them represent new genera and species, and are very
interesting, viz., Amphiute paulina, so far the first
instance of a calcareous sponge containing large, longitu-
dinally arranged, oxeote spicules both in the dermal and
gastral cortex, and Physcaphora decorticans, a tetracti-
nellid sponge, with a new type of microscleres. ‘There

SE FOP IEEE ETRE LNT IS

SST

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906 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

are also four new species, belonging to the genera Leu-
candra, Gellius, Raspaila and Pecillastra respectively.

Regarding the locality of the Sponges, it may be under-
stood, unless expressly stated to the contrary, that they
were all obtained from the neighbourhood of Sines, West
Coast of Portugal.

CALCAREA.

Order Homoca@.a.
Leucosolenia coriacea, Fleming. |
Several specimens of the usual character.

Order HETEROCGLA.

Leucandra aspera, O. Schmidt.

Von Lendenfeld (6, p. 125) in the year 1891 mentions
this species as ‘‘ beschrinkt auf das Mittelmeer.” Topsent
(9, p. 23) in the following year, describes it as from the
Azores. This is therefore only the second instance of —
this species being obtained outside the Mediterranean.

Leucandra bulbosa, n.sp.

Solitary, sessile, of bulb-like, or sometimes irregular,
shape, tapering upwards to a terminal osculum, which is
provided with a very small oscular frmge. Surface some-
what corrugated and hispid. The larger and more regular
specimens measuring about 22 mm. in diameter and 20
mm. in height. Width of osculum 2°5 mm. Colour (Gin
spirit) white or yellowish grey.

Canal system typical. Inhalant pores 0°04—0°07 mm.
in diameter, flagellated chambers 0°075 mm., exhalant —
canals 0°6 mm., or slightly more in diameter, sometimes
uniting in slit-like depressions. |

Skeleton: (1) Castral tetracts: apical ray 0:1 by 0°01
mm., basal ray 0°17—0°27 by 0°01 mm., oral rays 0.39—
0.45 by 0°01 mm. The basal and oral rays le in and
parallel to the gastral cortex, the apical ray projects at

SPONGES FROM PORTUGAL. 207 ©

right angles into the gastral cavity. The basal rays of
all the spicules are directed vertically downwards, the
lateral rays laterally and slightly upwards. (2) Triacts of
chamber layer: basal ray 0'4 by 0°022 mm.; oral rays
0°34 by 0°022 mm. (8) Dermal triacts: basal ray 0°2 by
0018 mm.; oral rays O17 by 0:018 mm. (4) Dermal
oxea, radially arranged and projecting, up to 14 by 0:075
mm. (5) Dermal club-shaped spicules, radially arranged,
only slightly projecting, 0°45—0°75 by 0°05 mm. (6)
Dermal hastates, minute, 0°07 by 0°002 mm. (7) Oscular
rhabds 0°42 by 0:0013 mm.

| Amphiute, n.g.

This genus belongs to Dendy’s family Heteropide
(Dendy, J, p. 75) as possessing a distinct and continuous
dermal cortex, covering the chamber layer and pierced by
inhalant pores, and also subdermal sagittal triacts. Its
flagellated chambers are sometimes elongated and radially
arranged, starting finger-like from large exhalant canals,
at other times quite irregular. Of the other three genera
belonging to this family (viz., Grantessa, v. Lendenfeld,
Heteropia, Carter, Vosmaeropsis, Dendy) Amphiute stands
nearest to Heteropia, as possessing large oxea, lying in
the dermal cortex, and arranged parallel to the long axis
of the sponge, but it differs from Heteropra in having a
similar layer of oxea in the gastral cortex. We find a —
strong resemblance to Amp/hiute in two genera belonging
to the Grantida, viz., Ute, O. Schmidt, with a layer of
longitudinally arranged oxea in the dermal cortex, and
Utella, Dendy, with a similar layer in the gastral cortex.
In proposing the name Amphiute in my preliminary
definition (4) of this genus, I had paid more attention to
these characters, than to the presence or absence of
subdermal triacts. If we accept Dendy’s classification,

,
h
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v
;
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i

208 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the presence of those triacts brings Amphiute under the
family Heteropide. Their absence would have shown
this genus to belong to the Grantide.

Diagnosis of Amphiute, n.g. The flagellated chambers
are sometimes elongated and radially arranged, sometimes
irregular. Dermal cortex and gastral cortex are both well
developed and both contain large oxea arranged parallel
to the long axis of the sponge.

Amphiute paulin, n.sp. (Pl. XIL., figs. 1—5; Pl. XTII.,

fon);

T'wo specimens were sent to me for examination. The
larger of the two is a colony of eight individuals, united
together at their bases (Pl. XII., fig. 1). The individuals
have a somewhat curved elongated cylindrical form,
tapering very slightly towards the distal osculum which
bears a small fringe. The larger individuals measure up
to 19 mm. in length and 3 mm. in diameter, the oscular
fringe is 1 mm.in length. The surface is smooth, and
shows a distinct longitudinal striation, due to the presence
of huge oxea in the dermal cortex. Colour (in spirit)
ereyish or whitish.

The canal system is sylleibid, and resembles closely
that of Vosmaeropsis macera, as described by Dendy (2,
p-. 182). The dermal pores are from 0°052 to 0°096 mm.
in diameter, and lead into inhalant canals which soon
become narrower. ‘The flagellated chambers are in many
cases elongated (0°44. by 0°'1 mm.) and open into very
wide exhalant canals, which narrow again before opening
into the gastral cavity. Thus the flagellated chambers
seem to be radially arranged less with respect to the

gastral cavity, than to the exhalant canals (Pl. XIII., fig. 1).
Sections, longitudinal or transverse, through the sponge |

show also a large number of spherical, oval, and irregular
chambers. But whether the chambers are really of those

SPONGES FROM PORTUGAL. 209

shapes, or whether this appearance is due more to the
direction of the section passing at different angles through
the chambers, is difficult to decide.

The skeleton is composed of seven kinds of spicules:
(1) Gastral tetracts, the facial rays measuring 0°132 to
0-16 mm. by 0°004 to 0°008 mm., the apical ray 0°056 to
0:076 mm. by 0:004 to 0°008 mm. (2) Subgastral triacts,
the basal ray 0°28 by 0°012 mm., the oral rays 0°092 to
O01 mm. by 0:012 mm. (8) Subdermal triacts, fewer in
number than the subgastral triacts, but of about the
same dimentions. (4) Dermal triacts, regular, each ray
01 to 012 mm. by00l mm. (5) Huge oxea, 1:2 to 2°5
mm. by 0°06 to 0°09 mm., occurring both in the gastral
and dermal cortex and arranged parallel to the long axis
of the sponge. (6) Rhabds, situated in the dermal cortex
and projecting at right angles, 0°2 by 0:0025 mm., or
longer. (7) Oscular rhabds; 1°2 to 2 mm., by 0'007 mm.,
forming a dense fringe.

Heteropegma nodus-gordu, Poléjaeff (7).

Represented in our collection by a single small colony,
easily recognised by the huge subdermal tetracts. Polé-
jaeff described this species first as from off the Bermudas
and Cape York.

SILICHA.

Order MONAXONIDA.

Halichondria, sp. ?

A single small encrusting specimen, 2 mm. in thickness,
yellowish-grey (in spirit), very soft and pulpy. Oxea
0°125 by 0°005 mm.

Reniera (cinerea, Grant ?).

Fistulous, consisting of three conical branches, the
largest of them being 22 mm. in height, 12 mm. in its
ereatest diameter, and the osculum 3 mm. in diameter.

910 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Very soft and elastic. Dermal skeleton unispiculous,
primary fibres of the choanosomal skeleton unispiculous,
sometimes bispiculous, secondary fibres unispiculous.
Oxea 0°088 by 0:005 mm.

Renera, sp.?

Small ridge-lhke specimen, 25 mm. in length, 8 mm. in
width, 8 mm. in height, with four large oscula along its
summit, the oscula 2 mm.in diameter. Colour (in spirit)
) almost black. Very soft, somewhat elastic. Skeleton

fibres unispiculous, rarely bispiculous. Oxea 0°084 by
0:0035 mm.

Renera, sp. ?

Small sessile specimen, 2 cm. in diameter, 5 mm. in
thickness, with three oscula each about 2 mm. in diameter.
Very soft and pulpy. Colour (in spirit) greyish brown.
Skeleton: meshes irregular, unispiculous. Oxea slender,
0-088 by 0:003 mm., sometimes stouter.

Remera, sp.?

About a dozen small finger-like specimens, attached to
the severed claw of a Crustacean, each about 12 mm. in
length, 1°55 mm. in thickness. Soft and elastic. Colour
(in spirit) brown. Oxea 0'104 by 0:005 mm.

Dactylochalina cylindracea, v. Lendenfeld (?).

The specimen in question offers a strong resemblance
in its external characters to the above species, as figured
by von Lendenfeld (5, Pl. II., fig. 1) in his Monograph
of the “‘ Horny Sponges,’ Pl. II., fig. 1. The sponge
is digitate, and consists of a number of slender cylindrical
branches arising from a common trunk. The entire height
of the specimen is 14 cm., the diameter of the branches 3
to4mm. The oscula are very small, less than 1 mm. in
diameter, and not raised. Consistency elastic. Colour
(in spirit) brown. The fibres are stout and contain a
large amount of spongin. The primary fibres, 0:045 mm.

:
:

SPONGES FROM PORTUGAL. A

in diameter, are multispiculous; the secondary fibres are
nearly as thick, but unispiculous. The oxea are stout,
0-108 by 0:009 mm. The dimensions of the spicules of
the type-specimen (from Australia) as given by von Len-
denfeld, are considerably less, and therefore I am in doubt
in regard to the identity of the two forms. Locality:
Lega.

Chalina, sp. ?

Three small finger-like specimens, about 20 mm. in
length, 2 mm. in thickness. Oscula very minute. Colour
(in spirit) yellowish-grey. With very little spongin.
Primary fibres multispiculous, about three spicules side
by side; secondary fibres.unispiculous. Oxea stout, 0°076
by 0°006 mm. Locality: Buarcos.

Chalina, sp. ?

Small, irregular specimen, encrusting Corallina. Prim-
ary fibres, with a considerable amount of spongin, uni-
spiculous, 0°024 mm. thick; secondary fibres very thin,
spongin scarcely covering the spicules. The spicules are
slender oxea, 0°076 by 0°003 mm.

EHuspongilla lacustris, Autt.

A single specimen, from a small river near Caldas de
Nixella, North Portugal.

Gellius pyrrht, n.sp.

This new species is represented by two fragments, the
larger of which is apparently a piece of a sessile, branching
specimen. It is 3°5 cm. in its greatest horizontal expan-
sion, 1 cm. in thickness, with about fifteen oscula which
are not raised, and are 1 mm. in diameter. Consistency
pretty firm and elastic. Colour (in spirit) greyish-yellow.
The skeleton consists of oxea which are very variable in
size, averaging 0°15 by 0°006 mm., and of sigmata which
are exceedingly slender, measuring 0°012 by 0:0004 mm.
or even thinner.

212 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Dendoryx imcrustans, Gray, v. viscosa, Topsent (9, p. 98).

The larger of the two pieces sent to me for examination ~
is roughly cylindrical, 5 cm. in height, 2 cm. in diameter,
and is apparently cut off from a much larger specimen.
The meandering ridges of its surface are very similar to
those of Dendoryx incrustans of the British Coast. The
strongyla of the ectosome have two minute spines at each
end, and measure 0:16 by 0:°005 mm. The spined styli
of the choanosome are 0°136 by 0:008 mm. The micros-
cleres are isochele, 0°016 mm. in length or less and
sigmata, 0°024 mm. in length.

Echinoclathria servata, Grant.

One specimen, of the usual character.

Raspailia formidabilis, n.sp.

One single specimen, 4 cm. in height, consisting of a
large number of bushy branches from a common trunk,
the trunk being 8 mm. in diameter. Entire surface of
the sponge exceedingly spiny, the spicules projecting a
good distance beyond its surface. Skeleton consisting of
two kinds of spicules, (1) smooth styl, straight or slightly
curved 1°5 by 0°02 mm., and (2) echinating spined styli,
0:095 by 0°008 mm.

Hymeniacidon carunculum, Bowerbank.

One specimen, massive, measuring 3°5 by 2°5 cm. hori-
zontally and 1°5 cm. in thickness, of the usual characters.

Tethya lyncurium, Lin.

A single small specimen, 1°5 cm. in diameter.

Order MONOCERATINA.

Huspongia (osculata, v. Lendenfeld ?). See v. Len-
denfeld (5). a
Represented by a piece apparently cut off from a large ©
specimen. Colour (in spirit) reddish-brown.
Aplysilla (archert, Higgin ?). See v. Lendenfeld (6).

ig:

SPONGES FROM PORTUGAL. FN

Represented only by a worn out fragment of the
skeleton.

Aplysinopsis, sp. ?

Two of the specimens are flat, sessile, not branching,
measuring 4 cm. horizontally and 7 mm. in thickness,
their surfaces being raised into blunt conuli. A third
specimen, possibly belonging to a different species of
Aplysinopsis, is apparently cut off from a larger specimen.
It consists of a basal portion with three branches and
measures 5 cm. in length, each branch being about 12 mm.
in thickness. Its surface is covered by sharp-pointed
conules, arranged in irregular ridges. Consistency elastic
and very tough. Colour (in spirit) yellowish.

Oligoceras collectriz, KF. EH. Schulze. See v. Lenden-

feld (4).
T'wo specimens, of a very crumbling consistency.
Hirama variabilis, F. EK. Schulze. See v. Lenden-
feld (5).

T'wo specimens, the larger of the two being flat and
sessile, measuring 5 cm. horizontally, 6 mm. in thickness.
Thickness of the fibres 0°07 to 0°12 mm., thickness of the
filaments 0°002 mm.

Order TETRACTINELLIDA.

Pecillastra armata, n.sp.

This species differs from all other known species of the
genus by possessing anatrizena in addition to other spicules.

Sponge massive, irregular, measuring 10 by 5 by 5 cm.
Surface even, rough to touch. Oscula 1 mm. in diameter,
scattered. Hxamuned in the dried condition.

The skeleton appears very confused, and is made up as
follows: Megascleres: (1) Oxea, of huge dimensions,
somewhat slantingly arranged towards the surface, straight,
or only slightly curved, measuring 3 by 0°055 mm. (2)

PEEL

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SS aoe SEL
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214 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Calthrops, the actines 0°45 by 0:045 mm. (8) Ortho-
triena, mostly deformed, rhabdome only slightly longer
than the cladi, of about the same dimensions as the
calthrops. (4) Anatrisvena, fewer in number than the other
megascleres, projecting beyond the surface of the sponge,
with wide and distinct axial canal in rhabdome and
cladi, the rhabdome measuring 2°5 by 0°02 mim., the

_ Cladi 0:09 by 0:02 mm.

The Microscleres are of two kinds, (1) Smooth Microxea
present in vast numbers and forming a felted mass
throughout the whole sponge, 0°17 by 0°0035 mm., (2)
Spiraster, 0°02 mm. in length.

Physcaphora decorticans, n.g. & sp. (Pl. XIIL., figs. 2 & 8).

The collection contains only a single fragment of this
highly interesting sponge, and that fragment apparently
represents only the ectosome of a tetractinellid sponge
which had become peeled off, as often happens in cases —
where the cortex is highly developed. The specimen is
a thin flat piece, of stony consistency, measuring hori-
zontally 4 by 8 cm. and 0°5 to 1 mm. in thickness. The
surface is raised in minute conuli which may bear the
oscula at their summits. Its colour (in spirit) is yellowish
white or greyish, here and there rusty, in some parts
almost transparent. ;

The skeleton consists of megascleres and microscleres.
The former are tylostyh, 0°51 by 0:008 mm., arranged in

bundles converging towards the surface of the sponge,

and raising it up in little conuli, and forming also a
support for the tissue round and below the oscula (PI.
XIII., fig. 3). The microscleres are of four kinds: (1) —

Spirasters, 0°014 mm. in length, forming a thin crust

along the surface of the sponge. (2) Spherasters, with -
large centres and very short rays, forming a layer just
below the Spirasters, 0°012 mm, in diameter. (38) Spher-

SPONGES FROM PORTUGAL. 915

asters, few in number, with small centres and long rays,

0°016 mm. in diameter, occurring here and there in the
lowermost part of the specimen. (4) “ Selenasters,”’
forming a thick layer below the spherasters with large
centres, in fact constituting the chief mass of the specimen.
These spicules correspond in structure and position to
the sterrasters of the genera Pachymatisma, Cydonium,
etc., and I have chosen the name from their faint resem-
blance to a half-moon. They are really more sausage-like
in shape and the generic name Physcaphora has been
adopted to express this. The full-grown spicule measures
0°08 by 0°028 mm., and fortunately a number of young
stages were met with, so that the development of the
spicule could be traced. In the youngest condition
present the spicules had the shape of rods, nearly straight
or slightly twisted, beset with minute spines (Pl. XIIL.,
fig. 2a). In the next stage the spicule is still pretty
straight, but the spines are large and numerous, although
still distinctly separated (Pl. XIII., fig. 2b). In the next
stage, the spicule has already its typical sausage-shape,
the spines are very closely set, but still recognisable in
their individuality (Pl. XIIL., fig. 2c). The last stage 1s
the full-grown selenaster, in which the spines, except
their most distal ends, are all fused so as to form one
solid mass (PI. XIII.,fig.2d). The distal ends of the spines
project a short distance beyond the surface of the spicule,
and being polygonal, chiefly hexagonal in transverse
section, offer a delicate pattern, when the spicule is being
focussed at different depths. A hilus is present as in the
Sterrasters. We thus see a great resemblance in the
structure and development of Sterrasters and Selenasters.
The chief difference is that in the Sterrasters all rays
start from a point, whilst in the Selenasters the rays
start from a line,

216 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. —

Regarding the systematic position of Physcaphora, we

have apparently to place it in the family Placospongide,

Sollas (8), provided we are right in our supposition that
the specimen in question is only the ectosome of a sponge,
and that the choanosome, when discovered, will not show
spicules of a different type. The Placospongide are
sterrastrose Tetractinellida, possessing, however, no te-
tract, but only monaxonid megascleres. Of the two
genera of this family, Physcaphora is more closely allied
to Placospongia, Gray, both possessing tylostyli, whilst
Antares, Sollas, possesses diactine spicules.

Thus we may perhaps propose the following diagnosis
of Physcaphora : Placospongide in which the megasclere
is tylostyle and in which the cortex is formed chiefly of
selenasters.

I now give a list of all the Sponges contained in the
collection. The classification is the same as used in my
‘ Revision of Bowerbank’s Nomenclature” (8).

CALCARHA.

Order HomMoc@La.
Leucosolema coriacea, Fleming.

Order HETEROCGLA.
Family Grantide : Leucandra aspera, O. Schmidt.
Leucandra bulbosa, n.sp.
Family Heteropide : Amplhiute paulim, n.g. & sp.
Family Amphoriscide: Heteropegma nodus-gordi,
Poleéjaeff.

SILICEA.

oe hee Order Monaxontpa.
. Family Haploscleride : Halichondria, sp. ?
: Reniera (cinerea, Grant ?)

SPONGES FROM PORTUGAL. -2°0 2.02 Q17

Reniera, sp.?
Renera, sp. ?
fienvera, sp. ? iG
Dactylochalina Hivindiees, v.
Lendenfeld (?).
Chalina, sp.?
Chalina, sp. ?
Huspongilla lacustris, Autt.
Gellius pyrrht, n.sp.
Family Peciloscleride: Dendoryx incrustans, Gray,
var. viscosa, 'T’.
Fichinoclathria seriata, Grant.
Raspaiha formidabilis, n.sp.
_ Family Azinellide : Hymeniacidon carunculum, B.
Family Tethyide: Tethya lyncuriwm, Lin.

Order MONOCERATINA.
Baily Spongide : Huspongia (osculata, v. Lendenfeld ?)
Aplysilla (archert, Higgins ?)
Aplysimmopsis, sp. ?
Oligoceras collectriz, F. E. Schulze,
Hirerma variabilis, F. EK. Schulze.

Order TETRACTINELLIDA.
Family Pachastrellide : Pecillastra armata, n.sp.
Family Placospongide : Physcaphora decorticans,
n.g, & sp.

LITERATURE.

1. Denpy, A. Synopsis of the Australian Calcarea
Heterocela, with a proposed classification of the
eroup and descriptions of some new genera and
species. Proc. R. Soc. Vict. (n.s.) V.,.pp. 69—116.

2. Dmnpy, A. Studies on the Comparative Anatomy of
Sponges. V. Observations on the structure and

218 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

classification of the Calcarea Heterocela. Quart.
J. Micr. Set. (n.s.), No. 138, pp. 15925

3. HanitscH, R. Revision of the generic nomenclature
and classification in Bowerbank’s ‘ British Spon-
gide.’. Trans. Liverpool Biol. Soc., VIII., pp.
173—206. it

4. HanitscyH, R. Amphiute, eme neue Gattung hetero-
coeler Kalkschwamme, Zool. Anz., XVII., p. 483.

5. LENDENFELD, R. von. A Monograph of the Horny

Sponges. London, 1889.

. 6. LENDENFELD, R. von. Die Spongien der Adria. I.

. Die Kalkschwamme, Zeitschr. wiss. Zool., LIII.,

. .. pp. 181—321, 361—483.

Re 7. PoLEJAEFF, N. Calcarea. Report on the. Scientific

| Results of the voyage of H.M.S. “ Challenger.”

i Vol. VIII. © | 3

| 8.. Sonnas, J. W. ‘Tetractinellida. Report of the Seine

. * tific Results of the voyage of H.M.S. “Challenger.”

o | Vol. XXV.

9. TopseNT, KE. Contribution a l’etude des Spongiaires

| del Atlantique Nord. In: Résultats des Campagnes

scientifiques, etc., Monaco, 1892.

EXPLANATION OF THE PLATES.

PuatTE XII.

Fig. 1. Amphiute paulini, n.g. and sp. Natural size.
{One or two individuals of the colony may be
somewhat misleading. The surface of the sponge
should appear smooth, not spiny, and the spicules
less pronounced.) ,

Fig. 9. Gastral tetract. The three straight rays are

- facial, the fourth, curved ray is apical. (x 150.)

ts

Fig.

Fig.
Fig.

Fig.

Fig.

Fig.

3.

SPONGES FROM PORTUGAL. 919

Represents both subgastral and subdermal
magens. (>< 150.)

Wermal triact;~ (< 150.)

Vertical section through the upper portion of
one of the individuals of Amphiute paulint. o.,
osculum ; g.c., gastral cavity. [In this figure as
well in fig. 1, Pl. XIII., Dendy’s plan has been
adopted to represent the collar cells diagram-
matically by red dots.] (xX 50.)

- Phare XIE.

Portion of a transverse section through one of
the individuals of Amphiute paulim. d.p, dermal
pore ; 2.c, inhalant canal; fc, flagellated chamber;
e.c, exhalant canal; g.c, gastral cavity. (x 80.)

Microscleres of Physcaphora decorticans,n.g. and
sp. a,b,c, young stages of Selenaster; d, adult
stage of the same. The somewhat eccentyrically
placed marking in ‘d’ is the hilus. /, various
forms of spiraster; e and g, forms of spheraster.
(x 500.)

Vertical section through the ectosome of
Physcaphora decorticans. o, osculum.

220

“NOTES on some points in the STRUCTURE of the
~CERATA of Dendronotus arborescens.

By Josepn A. Cuuss, B.Sc. (Vict.).

VICTORIA UNIVERSITY SCHOLAR IN ZOOLOGY ;
ASSISTANT-CURATOR OF THE DERBY MUSEUM, LIVERPOOL.

With Plates XIV. and XV.
[Read April 5th, 1895.] _

THE present paper 1s the outcome of part of the work
carried on in the Zoological Laboratory of University
College, Liverpool, under the direction of Prof. Herdman,
while in residence for the period covered by the Victoria
University Scholarship awarded me in June, 1894.

I. THE LIVER AND THE CERATA,

‘In the Second Report on the Nudibranchiata of the
L.M. B.C. District,* drawn up by Prof. Herdman and
myself, attention was directed to the published descriptions
by Alder and Hancockt and Dr. Rudolph Bergh{ of the
structure of the cerata of Dendronotus arborescens. These
distinguished zoologists described and figured the liver of
Dendronotus as giving off branched prolongations which
run upwards into the dorsal tentacles (rhinophores), and
other dorsal processes (cerata). Alder and Hancock
figure these hepatic ceca as conspicuous prolongations
from each side of the liver, while Bergh represents them —
as being of large size in the terminal twigs of the cerata,

* Trans, Biol. Secs, Vol. TLl., p.225.

+ Ray Society, ‘‘ British Nudibranchiata,” Part II., fam. 3, Pl. II.

+‘ Bijdragen tot de Dierkunde,” Natura artis magistra, Afl. XIII,
VIIL., p. 25, Amsterdam, 1886.

CERATA OF DENDRONOTUS. =tiy Oe

but does not figure the basal portions. As the result of
our investigations, both by dissections and by means of
thin serial sections we ventured to express our disbelief
in the existence of these hepatic ceca in the cerata; we
described and figured what we considered to be the
correct structure of the cerata, and showed that although
the liver is prolonged into short processes which run ‘in
the direction of some of the cerata, there is no hepatic
tissue to be found in the cerata themselves.* Last year
Dr. “Paul Pelseneer, in his ‘“ Recherches sur divers
Opisthobranches,’’+ in a paragraph on the digestive system
of Dendronotus briefly refers to our conclusions with
regard to the structure of the cerata. He says ‘‘ D’aprés
Herdman, le foie ne ramiferait pas dans les deux premiéres
pairs d’appendices dorsaux; les individus étudiés m’ont
présenté ces appendices pareils aux autres; le foie est
donc répandu dans toutes les saillies dorsales et dans le
corps, ou 11 occupe la méme position que dans Tritonia.”

It appears from this that Pelseneer has somewhat mis-
understood the results we then arrived at, for we not only
stated that the liver did not ramify into the first two
pairs of cerata, but also that it did not ramify into any.
Prof. Herdman at once suggested, as part of my work
while holding the University Scholarship, that I should
carefully revise all our previous investigations, examine
afresh our old preparations, and make what new ones I
should find necessary in order to settle definitely this
vexed question of the hepatic processes and dorsal cerata
of Dendronotus.

This I proceeded todo. I went carefully over the whole

* Loc. cit., p. 230.

+ Extrait du tome LIII. des Mémoires cowrownéds et Memoires des savantes
éranger, L’ Academie royale des sciences, des lettres et des beaux-arts de
Belgique, 1894. |

——

Sw ee es

'
ee

922 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of my series of sections, consisting of (1) a complete series
of transverse sections of the entire animal cut from end to
end, and arranged serially (containing some 600 or 700
sections), (2) a series of longitudinally cut sections, (3)
various sections of cerata isolated from the body, cut in
every direction. I minutely compared the figures published
in 1889, with the particular sections from which they were
drawn, and made fresh dissections of the animal. The
result of my work in this direction was to verify in every
particular the conclusions come to in our former paper
with regard to the cerata. I then set myself to consider
in what further directions I could proceed in order to
obtain additional evidence, and it occurred to me that a
series of sections cut horizontally, in a plane as nearly
parallel as possible with the creeping surface, would be of
great service. I could then pass from section to section,
tracing the liver from below upwards to its most dorsal
terminations, finally cutting transversely through the
bases of the cerata themselves; and by this means
determine exactly the relations of the liver to the bases of
the cerata. I obtained some living Dendronotus, from
Hilbre Island, and as the histological methods employed
were not described previously I will here describe them.
The Dendronotus is allowed to expand in a little
sea-water, and then deluged with sulpho-picric acid
(Kleinenberg’s formula), which by a rotatory movement
of the hand is made to whirl round in the vessel. This
treatment has the effect of fixing nudibranchs before they
can retract. Even with the more delicate species of
Eolide, which, with almost all other methods, break
away the cerata from the body, this method is usually
successful. The specimens are allowed to stand in this
fluid, changed once or twice, for two or three hours
according to size. They are then transferred to gradually

CERATA OF DENDRONOTUS. - JOS

increasing per centages of alcohol rising up to about 75%,
afterwards stained ‘‘in toto” in picrocarmine, treated
with acidulated alcohol, dehydrated, embedded in paraffin
in the usual way, and cut with the Cambridge “Rocking”
microtome.

- In order to make the sections of a manageable size, I
cut off the anterior and posterior moities, leaving the
central portion bearing the first and second pairs of cerata
only. A study of the sections showed me that they had
not been cut quite horizontal, the anterior end of the
section being relatively higher than the posterior. I
traced, first of all, the anterior prolongations from the main
lobe of the liver, which we had figured* running towards
the first pair of cerata. These I could follow through
several sections, lying alongside the stomach, to a point
immediately below the first pair of cerata. But I found
as I advanced through the sections and gradually rose to
a higher level, that the lver processes disappeared, and
disappeared before the stomach; which showed that they
terminated actually before the level of the top of the
stomach was reached. I then directed my attention to
the second pair of cerata. I picked up the main mass of
the liver at a point which I estimated to be immediately:
below them, and as I advanced through the sections,
eradually rising to a higher level, I traced the formation
of two lateral portions distinct from the central mass
(Pl. XIV., fig. 1, 2’). These lateral portions were evidently
the lateral caeca, previously figured} as running up towards
the bases of the second pair of cerata. But I now find
that these lateral masses actually disappear before the
central mass ; the right hand side one, two sections beyond
the one figured (fig. 1), and the left hand side one, four

o

ha c., Fl. X11., fig. 1.
quod c., Fl, XII., fig. 1.

224 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

sections beyond; while the central mass is continued for
twenty sections more. ,
- The section figured (Pl. XIV., fig. 1) shows the anterior
pair of cerata (c'), on the point of being separated off from
the body, while the second pair are not yet distinguishable
inany way. The liver (/) is here seen as I have described,
with a central mass and two lateral smaller masses, and
the ventricle of the heart (v) is shown lying in the peri-
cardial cavity (p c). one

Figure 2, 1s drawn from a section fifteen higher in the
series than fig. 1. Here the anterior cerata are quite
distinct, and the posterior (¢’’) are showing as distinct
prominences. The lateral processes of the liver (J’) seen
in fig. 1, have now disappeared, and the much reduced
i central mass remains. The coelomic cavity (c@) is also
much reduced, as the sections are now at a level at which
the mesodermal tissue of the dorsal body wall is appearing.
The distance of the liver mass from the place where the
cerata arise 1s very noticeable.

Figure 3 is a section thirty higher in the series than
fig. 2. Here the second pair of cerata (c’’) have now
become almost entirely separated off, although a slight
attachment still persists in one. The liver has entirely
disappeared some twenty sections before, but the second
pair of cerata have not yet become quite distinct even here.
The first pair of cerata are here (fig. 3) cut, among the
smaller terminal branches.

REET NETO

The minute examination of these sections of cerata,
whether taken before they are completely separated off
from the body and only showing as prominences at the
sides, or whether taken so as to cut through their base
just after separation, or whether sections of the terminal
twigs only, does not reveal any tissue that could possibly
be mistaken for liver. ;

-CERATA OF DENDRONOTUS. +e 995

Infact, I am convinced of the correctness of the
description given in our previous paper of the minute
structure of the cerata, notwithstanding the doubt thrown
upon it by Pelseneer. That description* is as follows :—
_ “1. Large spaces in the mesoderm, containing blood
corpuscles (Loc. cit., Pl. XII., figs. 2 and 3,¢s). These
run in the main longitudinally. They occasionally branch,
and they open into innumerable minute lacunew in the
mesodermal tissues, all of which here and there contain
blood corpuscles.

2. A good deal of pigmented connective tissue forming
branched masses and ramifying threads of a brownish
colour. These frequently, in a surface view of the
terminal branches of the cerata under a low power, give
rise to the appearance of a dark coloured granular central
eaecum such as that figured by Bergh (Loc. cit., Pl. IL.,
fig. 22). Sections, however, show the true nature of this
pigmented tract.

3. Masses of large distinctly nucleated cells lying in
meshes of fibrous connective tissue. These are possibly
mucus secreting glands. They occur chiefly in the
smaller meshes of the cerata.”’

I would respectfully suggest that a point of this
character, requires some of those “‘ complicated methods of
histological investigation’ which Pelseneer in his preface,
says he was unfortunately not in a position to make use
of. Without the aid of thin serial sections, I think it
would be impossible to determine with certainty this
point, and if, as was suggested in our previous paper,
these distinguished zoologists who differ from the con-
clusions come to, have worked without these aids to the
determination of minute structures, as I gather Pelseneer
has, then it is not surprising that they have been led
into error.

Loc, cit., p. 280.

226 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

It has been suggested that our Hilbre Island specimens
are of a variety differing in the structure of the cerata from
the specimens examined by others. I have endeavoured
to obtain specimens of Dendronotus from other localities
but without success. For myself, I do not think it likely.
However, I shall be very happy to exchange some of our
Hilbre Island specimens for some of these supposed
‘“‘ceratal hepatic caecal’’ bearing forms, if by so doing
Drs. Bergh and Pelseneer can be convinced of the correct-
ness of my conclusions, at least with regard to the Hilbre
Island Dendronotus.

A ie 0,

TI. Toe INNERVATION OF THE CERATA.

In the Quarterly Journal of Microscopical Science, a
| joint paper* was published by Prof. Herdman and myself,
Ei in which we showed from the examination of a series of
ti types of Nudibranchs, that instead of the cerata being
always innervated by the pleural ganglia, as Pelseneer
had previously supposed,t or always supplied by pedal
nerves, there are in fact various arrangements of the
nerve supply. We found from our investigations that
‘the dorsal lateral processes of the body wall, which we
call cerata, may be supplied entirely by the pleural
ganglia (e.g., Polycera and Ancula), or chiefly by the
pleural with a small supply from the pedal, by means of a
pleuro-pedal anastomosis (Dendronotus), or entirely by the
pedal ganglia (Tergwpes), or chiefly by the pedal ganglia
with a small independent accessory supply from the
pleural (as in Facelina).” If then the cerata are to be
regarded as homologous structures throughout the series
of nudibranchiata, the nerve supply cannot be taken as a

*“<QOn the Innervation of the Cerata of some Nudibranchiata,” Vol.

XXXIV., p. 541. |
+* Bulletin Scient.” ¢. XXIII, p. 489, Aug. 18th, 1891) ">=

CERATA OF DENDRONOTUS. 227

sure indication of homology, and we put forward the
suggestion that possibly the innervation has undergone
modification so that, while these ceratal outgrowths may
be truly epipodial, commencing as pedal structures,
supplied with nerves from the pedal ganglia, they may
have secondarily acquired in the different types mentioned,
as the result of changes in form, position and relation to
other organs, the various conditions of innervation which
we described. This, however, was merely a suggestion,
and requires further elucidation by the examination of the
nerve supply in other species, which as soon as I can
obtain the necessary material I hope to undertake.

The immediate object of this communication is with
regard to the nerve supply to the cerata of Dendronotus
arborescens. Since the previous paper in the Q. J. M. &.,
Prof. Pelseneer in his ‘‘ Recherches sur divers Opistho-
branchs”’’* refers to the conclusions we arrived at in
regard to this point. In Dendronotus we described and.
figured{ an anastomosis of a branch from the pedal nerve
with the pleural element of at least a part of the epipodial
nerve. We found from the examination of a complete
series of sections of an entire Dendronotus arranged
serially, that a nerve, arising from the pleural ganglia
(epipodial nerve), runs backwards for a short distance and
then divides into a dorsal branch (dorsal epipodial nerve),
and a ventral branch (lateral epipodial nerve) ; also that a
nerve arising from the dorsal aspect of the pedal ganglion
(dorsal pedal nerve) runs backward a short distance, and
then divides into two. The upper branch anastomoses
with the lateral epipodial nerve just after it has given rise

* Extrait du tome LILI. des Mémoires cowronnes et Mémoires des savantes
étranger. L’ Academie royale des sciences, des lettres et des beaux arts de
Belgique, 1894.

+ Loc. cit., Pl. XXXIV, fig..27.

298 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

to the dorsal epipodial nerve, so that the resulting nerve,

we pointed out, possesses both pedal and pleural elements.

Pelseneer while admitting the junction of the two nerves,
denies that the fibres pass from one to the other. In
a brief account of the nervous system of Phyllirhoe
bucephalum he describes and figures a branch from ‘‘le
nerf palléal’’ (epipodial nerve), which comes in contact
with a dorsal pedal nerve, and forms with it a plexus,
but with only a simple juxtaposition and without any
interchange of fibres. In a footnote* he says ‘‘ La meme
chose s’observe dans autres Tritoniens; par example
Dendronotus, ou Herdman et Clubb l’ont signalée, con-
cluant de ce fait que des fibres pédiewses passent dans le
nerf innervant les papilles dorsales. Or je me suis assuré
que chez Dendronotus, comme chez Phyllirhoe, il y a

simple juxtaposition locale des deux nerfs.” He again —

refers to it in similar language in a subsequent paragraph
dealing generally with the innervation of the dorsal
appendages.

It is obvious in a point like this in which the course of
the individual fibres of the nerve is concerned, that the
microscopic examination of thin serial sections with high

powers of the microscope is absolutely necessary, and that

it is impossible to determine it satisfactorily by means of
promiscuous sections or dissection only. It is somewhat
difficult to obtain sections in a perfect condition of
the region where this plexus is placed, owing to the
proximity of the odontophore, the radula of which is apt
to tear and destroy the sections, unless the animal has
been killed without undue contraction so that the odonto-
phore lies in its more normal position, in front of the

plexus. I have been fortunate enough, however, to ‘

obtain a series in which it is possible to trace the course
* Loc. cit., p. 42.

CERATA OF DENDRONOTUS. 299

of the nerves and their origin from the different ganglia.
With the higher powers of the microscope it is also
possible to make out the nerve fibres themselves, cut
transversely, in each of the sections. I have thus been
enabled to determine to my complete satisfaction that
there is a junction of nerves and not merely a running
alongside or juxtaposition without the fibres of the pedal
branch mingling with those of the pleural branch. In
order to demonstrate this I give drawings made with a
Zeiss Objective D from the various sections concerned in
this anastomosis of the two nerves. In Pl. XV., figs. 2,
3, and 4 are made from three consecutive sections, and
show the origin of the branch from the dorsal pedal nerve.
Fig. 3 is made from the section through the point of
origin; fig. 2 just anterior to it; and fig. 4 just posterior
to it. In these sections the transversely cut fibres with
the nerve sheath or perineurium can be distinctly seen.

Fig. 2 shows two nerves (a and 0b), lying in the coelomic
cavity (cw), with part of the wall of the cesophagus (@).
The nerve marked a@ is the epipodial nerve originating
from the pleural ganglia; the nerve marked 0 is the dorsal
pedal nerve arising from the dorsal aspect of the pedal
ganglia. For the sake of distinctness other structures in
the neighbourhood are left out.

Fig. 3, taken from the next section succeeding fig. 2,
shows a constriction appearing in 0, with the perineurium
or investing sheath extending inwards from the con-
striction, suggesting the beginning of the branch nerve ¢
which in the succeeding section, shown in fig. 4, is seen
separated by a complete sheath. Here although the
nerve b and its branch ¢ are not actually apart, still the
fibres are, and the succeeding four sections show this
branch gradually getting more distant from 0, and rising
towards a,

230 TRANSACTIONS. LIVERPOOL BIOLOGICAL SOCIETY.

In fig. 5, taken from the fourth section in the series —
after fig. 4, the branch c is seen some distance from 6 —
although not completely separated, there being a remnant
of connective tissue present. But it is much nearer the
nerve a and also connected with it by connective tissue. |

Fig. 6, taken from the next section, shows the pedal
branch c absolutely separated from 6, and in close
connection with a, although still surrounded by its own
sheath. But the nerve a is here seen to be giving rise
to a branch d. The next section (fig. 7) shows this
branch divided off by a complete sheath, so that here
there are sections across three nerves (a, d and ¢) which
are In organic connection, but the fibres of each are
completely separated by clear and distinct sheaths. They
remain in much the same relation to each other through
the following three sections. But in the next section
(fig. 8) a and ¢ come into closer relation, there is a
common investing sheath, while the portion of the peri-
neurium between the two has become thinner and less
pronounced; and in the next section (fig. 9) it breaks
down altogether and the two nerves unite, the fibres of
the branch ¢ becoming common, and enclosed in the
same sheath ag a. There are seen now but two nerves,
the upper (d), a branch from a, and the lower, formed of
the remainder of a plus the fibres of the nerve c, which —
is a branch from the dorsal pedal nerve 0. The nerve a
arises from the pleural ganglion and therefore contains
pleural elements, while ¢ is a branch of the pedal nerve },
which arises from the pedal ganglia, and contains pedal
elements, and therefore the resulting nerve (a+c) contains
both pleural and pedal elements. ne

The crucial point is evidently at the junction of a and
c, as these are the two nerves which Pelseneer admits
come into contact with each other, but the fibres of

OERATA OF DENDRONOTUS. ~~. -->. 281

which he says do not pass into the same sheath. I have,
therefore, examined with particular care the sections
concerned in this junction, and carefully noted any acces-
sory evidence with regard to the surroundings, shape,
conformation and other particulars seen in both sections
(figs. 8 and 9), and which help in any way to define exactly
the relations of the two nerves. There is a piece of con-
nective tissue, marked with an asterisk, in fig. 8 just
above the thin, but still distinct partition separating the
two nerves (a and c) at this point. In fig. 9 this piece
of connective tissue, of exactly the same shape and
relative position is present, and so marks exactly the point
where in the previous section the partition existed, but
which has here completely disappeared. There is no
doubt that the branch nerve (d), arising as it does from
a, looks, when viewed by dissection only, like the con-
tinuation of a, and that the nerve a+c. also when go
viewed looks like the continuation of c, and it is not
surprising that the mistake has been made of supposing
the junction merely a juxtaposition and not a union of
nerves. But when subjected by means of thin serial
sections to the higher powers of the microscope, it
obviates any chance of making errors of observation of
this character, and such a study of my sections convinces
me that the fibres of the branch ¢ pass into, and are
enclosed in the same sheath as a, and therefore that the
lateral epipodial nerve contains both pleural and pedal
elements, as originally stated by Prof. Herdman and
myself.

en

EXPLANATION OF PLATES.

Reference letters :—Pl. XIV., a, auricle; c’, first pair of
cerata; c’’, second pair of cerata; cw, colome; cs,

932 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,
ceratal sinus; dlv, dorso-lateral vein ; Ul, liver; 1', lateral —
lobes of liver; Js, blood lacune in mesoderm; m, muscle
bands; mt, muscular tissue; pe, pericardial cavity ; r,
rectum ; v, ventricle. }

PI, XV. a, nerve arising from the pleural ganglia
(epipodial nerve) ; 6, nerve arising from the dorsal surface
of the pedal ganglia (the dorsal pedal nerve); c, branch
nerve arising from the dorso- pedal nerve ; d, branch
nerve arising from the epipodial nerve; a-+c, the nerve
formed by the union of the epipodial with the branch c¢,
from the dorsal pedal nerve; ce@,'ccelome; @, cesophaeus ;
p.g., pedal ganglion; pl.g., pleural ganglion; p.n., pedal
nerve. |

PuaTe XIV.

‘Hig. il Horizontal section through the portion of the
-body of Dendronotus arborescens which bears
the first and second pairs of cerata, and passing
through the upper part of the visceral mass. —
The anterior end of the section is relatively
higher than the posterior so that the anterior
pair of cerata are seen almost separated off from
the body, but the posterior pair are not yet
distinguishable, the section passing below them,

‘Fig. 2. Taken from a section fifteen in number higher in
the series than fig. 1. The bases of the anterior
cerata are shown, cut through transversely and
completely separated from the body; and the
posterior pair are seen as distinct ee
at the sides. 3

Fig. 8. Taken from a section thirty higher in the series
than fig. 2. The upper parts of the first pair of
cerata are seen, cut transversely ; and the second
pair are cut through their bases and almost
completely separated from the body.

CERATA OF DENDRONOTUS. 933

Pratt XV.

Fig. 1. Diagrammatic scheme of the anterior part of the
dorsal and lateral epipodial nerves of Dendronotus
arborescens, showing in lateral view their origin
from the ganglia, their branches, and the anas-
tomosis between a and c.

Fig. 2. Drawn from a T.S. section (the 283rd in the serial

series) through the body of Dendronotus just
anterior to the point of origin of the branch from
the dorsal pedal nerve (in the plane marked by
the dotted line 2 in the diagrammatic scheme,
fig. 1). The relative position of the nerve

a (epipodial nerve) arising from the pleural
ganglion, of the nerve 0 (dorsal pedal nerve)
arising from the pedal ganglion and of the
cesophacus @ is seen.

Fig. 3. Drawn from the next section behind fig. 2 (in the
plane of the dotted line 3 in fig. 1). Here the
branch c. is beginning, and the perineurium is
seen partly cutting it off from 6b (dorsal pedal
nerve). |

Fig. 4. Drawn from the next section posterior to fig. 3 (in
the plane of the dotted line 4 in fig. 1). The
branch ¢ is now completely invested in its distinct
sheath, and its fibres are completely separated
from those of 6 (dorsal pedal nerve).

Fig. 5. Drawn from the fourth section posterior to fig. 4
(in the plane of the dotted line 5 in fig. 1). The
branch ¢ is now seen some distance from the
pedal nerve b, and close to the epipodial nerve «.

Fig. 6. Drawn from the succeeding section to fig. 5 (in the

plane of the dotted line 6 in fig. 1). The epipodial
nerve a is here seen to have a distinct constriction,
with a part d being separated off,

ies i ne from the next section to ion 6 is in ‘the
plane of the dotted line 7 in fig. 1). Shows the
portion d, completely separated by a dis

sheath, so that there is a group of three a
cut transversely, united by connective tissue, v2z.,
@ epipopdial nerve proper, c the branch from the
pedal nerve, and d the branch from a. — ae

Fig. 8. Drawn from the fourth section behind fig. 7 Gas P
the plane of the dotted line 8 in fig. 1). Shows —
the three nerves a, d, and c still surrounded by —

Fig. 9. Drawn from the succeeding section to fig. 8 (
the plane of the dotted line 9 in fig. 1) Sho
that the partition separating a and ¢ has bro

down and the fibres of the two nerves are
invested in the same sheath. a <a

235

[WORK FROM THE PORT ERIN BIOLOGICAL STATION. ]

NOTES on SOME SPECIMENS of Synapta
inherens from PORT ERIN.

By Herpert C. CHaDwick.
With Plates XVI. and XVII.
[Read January 11th, 1895.]

DURING a visit to the Port Erin Biological Station in
September, 1893, Prof. Herdman drew my attention to a
number of living specimens of Synapta inherens, which
he had found in the muddy eravel at about half-tide mark
immediately below the station. As the species had not
been identified, I gladly assented to the suggestion that I
should examine and report upon the specimens. A
further supply was obtained, and after being kept in clear
sea-water for some days, in order that the contents of
their intestines might be voided, they were fixed in an
extended condition by the addition of sulphuric ether to
the sea-water in a tube, and finally preserved in 70 %
alcohol. Since the time above mentioned I have carefully
examined the specimens; and I would like at this point
- to acknowledge my indebtedness to my friend Dr. C. H.
Hurst of Owens College, for placing at my disposal some
large specimens of the closely allied species Synapta
digitata, from Naples, by the dissection of which I have
been able to supplement the results obtained from dissec-
tion of the smaller and less easily examined British
specimens. My thanks are also due to my friend Mr. F.
W. Gamble, M.Sc., of Owens College, for valuable assist-
ance with the literature.

Specimens for sectioning were prepared by decalcifica-

9236 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

tion in a 10 % solution of nitric acid, and staining in a
rather weak solution of borax carmine, the specimens
remaining in the solution for from two to three days.

I think it advisable to preface my description of the
specimens from Port Erin by stating the characters of the
genus, and those by which the three British species are
distinguished by Prof. Bell in his ‘‘ Catalogue of British
Echinoderms in the British Museum,” published in 1892.
Genus Synapta: Tentacles ten to twenty-five; deposits
anchors, anchor plates, and fine granules ; hermaphrodite.

Synapta inherens. ‘Twelve tentacles, each with six or
seven digitate processes on either side ;* a single madre-
poric canal; one to three Polian vesicles; edge of fluke
of anchor serrated; several of the anchor plates with
serrated holes; miliary granules not largely developed.

Synapta buskt. Eleven tentacles, with three true
digits and two or three transverse .ridges which look like
digits; a single madreporic canal; one Polian vesicle.
Anchor plates with seven dentate holes, and long narrow
handle. ;

Synapta digitata. ‘Twelve tentacles, each with four or
five digitate processes on either side; a single madreporic
canal and Polian vesicle. Fluke of anchor smooth or
slightly serrated; the holes of the anchor-plates smooth
or with fainter serrations than in S. wmherens; miliary
granules numerous and in places crowded.

Several inconsistencies between these definitions and ~~

the figures which adorn the plates of the Catalogue impair
the usefulness of both. S. inh@rens is said to have
‘“ several’ of the anchor plates with serrated holes; while
each of the five plates figured have serrated holes. S.

-* Prof. Theéel, in his Report on the Holothuroidea of the ‘‘ Challenger” |
Expedition gives the number of digitate processes on the tentacles of S. inhe-
rens as from twelve to fifteen.

NOTES ON SYNAPTA. , 937

buski is described as having anchor-plates with ‘‘ seven
dentate holes and long narrow handle.” ‘The three figures
represent plates with five or six large holes and a number
of smaller ones, all of which have perfectly plain margins,
the handle of the anchor being short and stout.

S. digitata is said to have tentacles ‘‘ with four or five
digitate processes on either side,’ while the figure on
Plate VII. of the Catalogue represents a circlet of tenta-
cles with two digitate processes only on either side. This
figure is a fairly accurate representation of the tentacles
of a Synapta from Millport, given to me by Mr. P. Cameron
of Manchester, and of the tentacles of the large specimens
labelled S. digitata from Naples, which were entrusted
to me by Dr. Hurst. Of the three forms just enumerated
S. digitata appears to me to be distinct, but I am strongly
inclined to think that were specimens from a sufficient
number of localities available for examination the other
two would be found to be inseparable.

I have referred the specimens from Port Erin to Syn-
apta inherens. It is true that the number of digitate
processes on the sides of the tentacles never exceeds four,
and is very commonly three (Pl. XVIL., fig. 1). The
flukes of the anchors are, however, in nearly all cases
distinctly serrated, and not more than ten per cent. of the
anchor plates have holes with plain margins (Pl. XVIL.,
fig. 6), the denticles being well marked, though not so
numerous as those of the plates figured by Prof. Bell.
The so-called miliary granules, though numerous, are not
so thickly scattered as are those of the large specimens of
S. digitata from Naples. There is but one Polian vesicle,
which attains a very considerable length (Pl. XVL., fig.
1, p.v.) The lateral borders of the twelve tentacles are
crowded with calcareous spicules of minute size and
simple form, most of them being dumb-bell shaped. As

238 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the peristomial disk is approached, the shape gradually
changes until it becomes distinctly lenticular.

Upon the oral face of each tentacle are seated in two
irregular rows from two to eight peculiar bodies which
have been described as tentacular pedicels (Pl. XVIL., fig.
1, tp.) In shape they resemble a wine glass without the
stem (fig. 8, a), and so suggestive of a sucker is their
appearance when seen in situ, that the older zoologists
may well be excused for comparing them with the tube-
feet of Asterids and Hchinoids. In minute structure,
however, they are entirely different. Each consists of an
outer envelope composed of ordinary ectoderm cells (fig.
3, 6), enclosing a cup-shaped mass of cells, also ectodermal,
but of a more specialised kind. They are much longer
than ordinary ectoderm cells, especially those which lie
at the bottom of the cup, and the nucleus is basal. It is
highly probable that the free ends of these cells bear long
cilia, but I have not satisfied myself upon this point.
Occupying the same position on the tentacles of S. digi-
tata are a number of bodies (sixteen to twenty-four), first
described by Quatrefages as suckers, and since, by Hamann
as sense organs, which are obviously similar in structure
to those just described (Pl. XVII., fig. 2). There are,
however, certain important points of difference. The
mass of specialised cells does not project above the surface
of the tentacle, and the cavity which they surround in
S. inherens is here occupied by a solid core of elongated
cells, the basal ends of which pass into a bundle of nerve
fibres which can be traced into continuity with the ten-
tacular nerve. Hamann states that the free ends of these
cells are ciliated. I have examined my sections of S.
inherens with a view of discovering a nervous connection
if such existed, between the organs I have described and
the tentacular nerves, but without SUCCESS.

ia)
vw

a a
cs
7

NOTES ON SYNAPTA. 239

The body walls conform to the typical structure (Pl.
XVI., figs. 1 and 2). Beneath the ectoderm (ect) is a
thick layer of connective tissue (ctl). Then comes the
layer of circular muscular fibres (cm) lined by the ciliated
epithelium of the body cavity. |

The bodies to which I have twice referred under the
name of miliary granules are ectodermal structures which
project wart-like above the general surface of the body
(Pl. XVI., figs. 1 and 2, mt). In the substance of each
is lodged a relatively large ganglion-like mass of nerve
tissue (Pl. XVIL., fig. 7). From the latter a large number
of fibrillar processes radiate, some if not all of which
become continuous with the filiform basal ends of the
ectoderm cells, the ‘‘ sense cells’’ of Hamann and other
writers. Many of the ganglion-like masses are continuous
with bundles of nerve fibres which most probably arise
from the radial nerves. Another point of interest is the
presence in the connective tissue layer of what for want
of a better name, I have called giant cells, in allusion to
their large size. Occasionally they occur in groups of two
or three, but as a rule singly (Pl. XVII., fig. 8). The.
protoplasm is finely granular, and the majority of the
cells are multinuclear, the larger ones invariably so. They
seem to occur only in the most superficial portion of the
connective tissue layer, just beneath the ectoderm.

The nervous system consists of a circum-oral ring
(Pl. XVL., fig. 1, conr) from which five radial nerves (77)
proceed along the radu of the body. A well marked sheet
of nerve fibres is given off from the circum-oral ring to
the oral face of each of the tentacles (tn/). Seen in trans-
verse section this presents a crescentic figure, lying just
beneath the ectoderm cells (Pl. XVIL., fig. 5). The cir-
cumoral ring almost completely encloses a tubular space
(ts). - This, like the nerve ring gives off branches which

240 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

lie beneath and are partly bounded by the radial nerves.
The circular water vessel (cwv) gives off branches (twv)
to each of the tentacles, entrance to the tentacle being
made on the inner side of the calcareous ring (cr). Other
appendages of the circular water vessel are the madreporic
tube, or water tube, (w) and the single Polian vesicle (pv).
The former, a simple tube lined by (probably ciliated) epithe-
lum, is seated on the upper surface of the water vessel, and
its free end opens into the body cavity; the latter is also
tubular, and of great length, but its free end is imper-
forate. It depends from the lower face of the water vessel.
The intestine (7) though much sacculated and occasionally
bent upon itself, runs a practically straight course from
mouth to anus, being suspended from the dorsal interra-
dius by the mesentery (m). It is accompanied in its
course by the typical dorsal and ventral vessels (dv and vv).
The genital organ consists of two bundles of cecal tubes
(g) which lie one on either side of the mesenterial fold
which suspends the intestine. The main tube of the left
bundle joins that of the right by penetrating the mesentery.
A single duct is thus formed which, passing forwards in
close proximity to the water-tube opens by a single pore
between two, and just outside of, the ring of tentacles
(ga). Attached to the fold of mesentery by which the
intestine is suspended, just at the point where it joins
the body wall, there occur a great number of funnel-shaped
bodies with strongly everted margins, and lined by ciliated
epithelium (Pl. XVI., fig. 1, cf). These are the ‘‘ ciliated
funnels” of Semper. Hach funnel is attached by its
pointed apex to the mesentery ; and in addition to those
borne by the intestinal mesentery, another series, appear-
ing to the naked eye as an irregular but quite continuous
line, occupies the middle of the interradius next on the

Py
= t

a a

_ — —_—" =

ee ee ee a ee a ee ee ae ee a ee ee 2

NOTES ON SYNAPTA. QAT

left of the dorsal one. A few also occur scattered over
the walls of the colom. Their function is at present
unknown.

EXPLANATION OF Pirates XVI. and XVII.
List of reference letters.

of Ciliated funnels; cm circular muscles; conr circum
oral nerve ring; ctl connective tissue layer of body wall;
cwv circum-oral water vessel; cr calcareous ring; dv
dorsal vessel; e ectoderm; g gonad; ga genital aperture;
zintestine ; /mb longitudinal muscle bands ; m mesentery;
mlt muscular layer of tentacles; mt miliary granules;
pv Polian vesicle; 7m radial nerve; ¢ tentacles; tp ten-
tacular pedicels; tnd tentacular nerve; ¢s tubular space
underlying nerve; vv ventral vessel; w water-tube.

Pruate XVI.

Fig. 1. Diagram of the anatomy of Synapta iherens,
constructed from dissections and serial sections.

Fig. 2. Diagram of a transverse section of Synapta
imherens.

PLATE XVII.

Fig. 1. A tentacle of Synapta inherens showing the so-
called tentacular pedicels in situ.

Fig. 2. Outline figure of a tentacle of Synapta digitata,
showing the bodies described by Hamann as
sense organs.

Fig. 8. (a) A tentacular pedicel viewed in situ; (0)
vertical section of the same.

Fig. 4. Diagram of a longitudinal section of a tentacle of
Synapta mherens.

7"
. <eo eae

242 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

igs oe Transverse section of a tentacle of Synapta Be
inherens, taken at the level of the dotted line*
in fig. 4. | | Aus
Fig. 6. Plates and anchors of Synapta inherens; (a) is an 5 iy
abnormal form of anchor; (6) smaller calcareous er
plates. |
Fig. 7. Vertical sections of three miliary granules of —
Synapta inherens, showing the ganglion-like
nerve endings.
Fig. 8. Section of ectoderm and connective tissue layer F
of the body wall, showing giant cells in the latter. aa

» tet

243

[WORK FROM THE PORT ERIN BIOLOGICAL STATION. ]

moe Okh on the MEDUSA of the L.M.B.C.
DISTRICT.

By Epwarp T. Brownz, B.A.

UNIVERSITY COLLEGE, LONDON.

[Read March 8th, 1895.]

Ir is nearly nine years since the first report on the
Medusz of the L.M.B.C. district was published by Mr.
J. A. Clubb (7). It contains a list of medusz collected
by Prof. Herdman off the south end of the Isle of Man
in 1885, and by Byerley (6) and other naturalists who
have worked in Liverpool Bay. Most of these species I
have recently taken at Port Erin, some others, which I
have not met with and which are mentioned in the first
report, are again recorded in this report so as to make the
present list as complete as possible.

My own records are chiefly based upon work done at
the Port Erin Biological Station between April 28th and
June 7th, 1893, and also between March 22nd and May
14th, 1894. I have confined my attention to the small
Hydromedusze and have not specially searched for the
other groups. Mr. W. 1. Beaumont (5) in 1893 published
an account of the Lucernarians of Port Erin. I owe to
Mr. Beaumont my sincere thanks for the great aid which
he has so often generously given in helping me to collect
specimens of the Hydromeduse. I have not met with
any new species, but have added a few interesting ones
to the list of Manx medusz and also show that some,
described as species, are stages in development of other
species already recorded.

944. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

This report can only be regarded as a preliminary one,
| as it is simply an abstract of some of my notes. I hope
| . at a later date to publish an account of each species with
| figures. The great obstacle which prevents progress is
. not being able to collect sufficient specimens of a species:
so as to trace its development and to see how far it is
necessary to allow for variation. So far my attempts to
rear medusze from their hydroids, or to keep them in
confinement have been failures. The rearing of a medusa
in confinement is the only safe method for tracing its
life-history, and until this can be done the uncertain
method of catching a sufficient quantity of specimens
must be adopted. Some species have a characteristic
feature, like the eight radial canals of Melicertidiwm,
which distinguish them from other allied species, and by
which it is easy to identify them at any stage in develop-
ment. But unfortunately the majority of species have
not such characteristic features, and it is difficult to
- distinguish the early stages of different species. The old
| genus T’hawmantias of Forbes, which has been divided
{

into many families and genera by Heckel, still remains a
ereat stumbling-block to the medusa-collector. Another
difficulty which has yet to be overcome is the final indenti-
fication of a number of doubtful species of hydroids, which
are known to bud off meduse. When the life-histories
of the hydroids with their meduse are known and the
specific differences correctly determined there will be no
difficulty in adopting a single system of classification.
At present two systems exist. one for the hydroids and
another for the meduse. I have used in this report
Heckel’s system of classification and the names of his
genera, but have also given the principal synonyms and
references to those species which are mentioned by Forbes,
Hincks and Allman. There is no need in this kind of

Daas
——
_— SST
ELE AGES ESO ION SEALE II

ee

— a
Se eran alae aaa

a
5 p

tl el DE 2 tae

L.M.B.C. MEDUSA. DA5

report to give a description of the species. Nearly every
species mentioned in this report is given by Forbes in his
monograph.

My latest method for catching medusez is to use a long
tow-net made of bolting silk with a mesh not exceeding
half a millimetre. At the end of the net is fastened a
zinc can, instead of the usual glass bottle which generally
breaks when the contents are specially wanted. I prefer
to use a small rowing boat and work very gently against
the tide with just a slight pull on the net. If the
meduse are not visible at the surface, I sink the net with
a weight about two fathoms, and if not successful, try
near the bottom. The net is left down from 10 to 20
minutes and the contents emptied into a large glass-bottle.
It is best to pick out at once some of the best specimens
with a large pipette and place them in another bottle full
of clean sea-water. Meduse are very delicate, and
quickly die if crowded together in a small bottle.

The following is one of the best methods for killing
and preserving small meduse. Place the medusa in a
large watch-glass with sufficient water to float in. Add
with a pipette about 5 drops of a 2 per cent. solution of
cocaine, and about 10 minutes later add another 5 to 10
drops; the quantity depends upon the species and its
activity. When the medusa has become motionless and
the tentacles expanded, add suddenly a saturated solution
of picric acid. It is best to reduce with a pipette the
quantity of water in the watch-glass just before adding
the picric acid, and use plenty of the latter. Often when
the picric acid is added a precipitate of cocaine is thrown
down which must be removed at once by a pipette and
fresh picric acid added. Specimens may be left in picric
acid about half an hour and gradually transferred into 80
p-c. alcohol. I strongly recommend the use of the best

246 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

glass-stoppered bottles and perfectly pure spirit for storing
specimens in. ,

Finally I thank Prof. Herdman and the Committee for
the use of the Port Erin Biological Station.

HYDROMEDUSZ.
Order ANTHOMEDUSAK.

Family CoDONIDZ.

Codonium pulchellum, (Forbes 16).
Sarsia pulchella, Forbes (9).
Syncoryne pulchella, Allman (4).

This species was fairly abundant during May and the
early part of June, 1893, in Port Erin Bay. The bulb at
the base of the tentacles varies in colour in different
individuals. It is usually of a brownish colour, but in
some specimens either orange, crimson, or greenish. The
“ Scheitel-Aufsatz’’—(the prolongation of the manubrium
into the substance of the umbrella, forming a kind of
apical knob to the manubrium) is present in all specimens;
but the ‘ Stielcanal’’ (which runs from the apical knob
to the exumbrella, is not a true canal or tube, but the
remains of the original canal connecting the young medusa
with its hydroid) in some specimens is very narrow and
difficult to see. In a few specimens it is entirely absent.
Heckel divides the genus Sarsia of Forbes into two genera,
Codoniwm and Sarsia. The species of the former genus
are characterized by the possession of a ‘‘ Scheitel-Aufsatz”’
and a ‘‘Stielcanal.’”’ The genus Sarsia of Forbes still
requires careful revision.

Codonium pulchellum swims actively about an aquarium,
and usually has its manubrium and tentacles expanded.

It resembles Sarsia tubulosa, Sars, rather than S. pul-
- chella in its habits. The manubrium and tentacles, when

L.M.B.C. MEDUSA. DAT

fully expanded, are about twice the length of the umbrella.
The umbrella of the smallest specimen taken measured
4 mm. in length and 3 mm. in width: the largest
specimen 9 mm. long and 6 mm. wide. Other specimens
-measured—6 mm. long and 6 mm. wide; 6 mm. long and
7mm. wide; 8 mm. long and 8 mm. wide.

Sarsia pulchella may be an early stage of S. tubulosa.

Distribution.—Shetland, St. Andrews, Plymouth, Firth
of Clyde.

Sarsia tubulosa, Sars, (9).

Byerley (6) records this species.—‘‘ Caught in the
Mersey, rare, Mr. Price.” Haddon (4) records S.
tubulosa in Dublin Bay.

I have taken in Port Erin Bay several specimens of
young Sarsia, but have not been able to determine for
certain the species.

Dipurena halterata, (Forbes 15).

— Slabberia halterata, Forbes (9).

Two specimens taken in August, 1892. One measured
91 mm. in length and 2 mm. in width. Manubrium
extended to 8 mm. Tentacles 2 mm. long. No repro-
ductive organs present. The medusa may easily be
distinguished by the large knob, containing nematocysts,
at the free end of each of the four tentacles.

Distribution.—Jersey, Mount’s Bay (Cornwall), Ply-
mouth,

Steenstrupia rubra, Forbes (9).
Steenstrupia galanthus, Heeckel (15).
Steenstrupia flaveola, Forbes (9).
Forbes describes and figures the above species, the
former taken off the Shetlands, the latter at Penzance.
They mainly differ in colour, Heckel regards them as

2948 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the same species. I took two specimens off Port Erin in
August, 1892. The contents of the tow-nets were unfor-
tunately preserved at sea, so that the colour of the medusa
was destroyed. In shape and size they resemble the
figures given by Forbes of Steenstrupra. Heckel regards
Steenstrupia galanthus as the medusa of the hydroid
Corymorpha nutans, of Allman and Hincks, but not of
Sars. The various species of Corymorpha and their
meduse require careful revision.

Huphysa aurata, Forbes (9).

A single specimen of this medusa was taken on May
5th, 10th, and 20th; and five specimens on June 5th,
1893.

The umbrella of the largest specimen measured 4
mm. in length and 3} mm. in width; the smallest 34 mm.

long and 2mm. wide. One specimen taken on June 5th ~

with ring-canal coloured in several places with a crimson
pigment. The three bulbs, without tentacles, had their

centres coloured with a brilliant crimson pigment, but

the basal bulb of the large tentacle was very faintly

- coloured with crimson.

One specimen, taken on March 22nd, 1894, the first
day of tow-netting, measured 38; mm. in length and
2mm.in width. None seen until April 18th when three
were captured, the largest 6 mm. in length and 43 mm.
in width. After this date a few were regularly taken 1 in
the tow-net.

The umbrella is always longer than wide, and laterally
very thick; the latter a characteristic feature by which
this species may be readily distinguished from other
uni-tentacular meduse. The exumbrella has a few
scattered nematocysts.

Forbes (9) in his description of this species writes :— _

‘The orifice (of the umbrella cavity) is rather contracted

avin Bee MTEL wer) of DAY

and square. At each of the four angles is a large
diamond-shaped ocellus, the upper half of which is
bright golden yellow, and the lower half vivid scarlet or
crimson. From each of the ocelli springs a short, reflexed,
cylindrical, yellow tentacle, which I have never seen to
extend itself. From one of the ocelli, below the short
tentacle, arises a long and thick one, highly extensile, and
of a golden colour, usually presenting a club-like shape.”

This statement does not give a quite correct description
of the species. I have examined the tentacle bulbs of
about two dozen specimens and failed to find an ocellus
definite in shape and colour. The tentacle-bulbs are
similar in shape, but one has a large tentacle and the
other three are without tentacles. Forbes describes and

- figures each tentacle-bulb with a minute, reflexed, yellow

tentacle, and one bulb having an additional large tentacle.
The ‘“‘ minute tentacle’”’ is the lower part of the tentacle-
bulb, which slightly curls over the margin of the umbrella
to which it is fixed.

The colour of the tentacle-bulbs varies considerably in
different specimens. A few had no colour whatever in
the bulbs and only the large tentacle faintly coloured
with yellow. One specimen had the three bulbs, without
tentacles, quite colourless, but the tentacle-bulb of the
large tentacle of a yellowish colour. Most specimens,
however, have yellow or orange pigments in the lower
part of the bulbs, the part which corresponds to ‘‘ minute
yellow tentacles” of Forbes. In the most brillantly
coloured specimens the interior of the tentacle-bulbs has
a crimson pigment, the part which corresponds to the
crimson ocellus of Forbes. In one specimen the crimson
colour extended a little way along the ring-canal on each
side of the bulbs, and in two other specimens, brilliantly
coloured, the whole ring canal.was crimson. Crawford

250 TRANSACTIONS LIVERPOOL BIULOGICAL SOCIETY.

(8) has also observed a crimson ring-canal in Se
taken at St. Andrews.

Alder, (8) describes and figures the medusa of Core
morpha nana, Alder, (= Halatractus nanus, Allman 4),
which he bred from the hydroid. Heeckel (15) considers,
from Alder’s description of this medusa and from the
description of Huphysa aurata by Forbes, that the
medusa of Corymorpha nana and EL. aurata are identical
and gives in his ‘‘ System der Medusen” Halatractus
nanus as the hydroid of the medusa Huphysa aurata. I
believe Alder is the only person who has ever taken
Corymorpha nana; and Heckel apparently has not met
with Huphysa aurata. The figure given by Alder of the
medusa of Corymorpha nana has not the specific charac-
ters of H. aurata. The umbrella of Corymorpha is about

as long as wide, and thin; the umbrella cavity is conse-

quently very large. The umbrella of Huphysa is always
longer than broad, and laterally very thick; also about one
third of the aboral end is formed of solid tissue; the
umbrella-cavity is small compared with the external size
of the umbrella. The manubrium of Corymorpha is long

and extends beyond the velum, but the manubrium of

Euphysa is about two-thirds the length of the umbrella-
cavity and has never been seen to extend beyond the
velum. Huphysa aurata may be easily recognised by
the shape and great thickness of the umbrella. I think
that the meduse of Corymorpha nana and Huphysa
aurata do not belong to the same species, and that the
hydroid form, if one exists, of Huphysa has yet to be
discovered.

Heeckel (15) describes and figures in his mage
another species of Huphysa—EH. mediterranea—which he
obtained at Villafranca. The umbrella is longer than

wide (8 mm. long, 2 mm. wide), and thick. Three tenta-

yy"

L.M.B.C. MEDUSA. Fe QO E

cle-bulbs, without tentacles, golden yellow, with purplish
red ocelli. The other tentacle-bulb, golden yellow, bearing

- a large golden red tentacle, without a purplish red ocellus.

Ring-canal purplish red. When I compared a living

specimen of H. awrata, which possessed a crimson ring-

canal and tentacle-bulbs coloured with orange and crimson,
with the coloured figures given by Heeckel in his mono-
eraph, I failed to find sufficient differences to separate
H. aurata from EH. mediterranea. |

Heckel makes the chief specific difference the absence
of the purplish red ocellus in the tentacle-bulb of the large
tentacle. In H. awrata there is no proof that the part of
the bulb coloured crimson is really an ocellus. In some
specimens the colour is absent, but present in others in
different shades of crimson. One specimen, taken in
1893, had the three bulbs, without tentacles, conspicuously
coloured with crimson, but the bulb at the base of the
large tentacle had only a faint tinge of a crimson colour,
scarcely visible with the microscope. I have noticed in
other species of meduse a great variation in colour.
Amphicodon fritillaria has usually dark brown or reddish

brown tentacle-bulbs, and all the canals colourless; but

in a few specimens the tentacle-bulbs are of a brilliant
red colour and also the radial canals. |
Distribution.—Shetland, St. Andrews.

Amphicodon fritillaria, (Steenstrup 15).
Coryne fritillaria, Steenstrup (21).
Steenstrupia globosa, Sars (20).
Amphicodon globosus, Heeckel (15).
Diplonema islandica, Green (18).
Steenstrupia owen, Green (18).
Hybocodon prolifer, Agassiz (2) Heckel (15).
Amphicodon amphipleurus, Heeckel (15).

Pe

ERT

. medusa which he described as follows :—Umbrella bell-

t

252 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.
Fs Steenstrup (21) during April and May, 1840, off Reike-

wig, Iceland, obtained specimens of a free-swimming

shaped, little longer than broad. Manubrium about 2 the
length of the umbrella cavity. Margin of the umbrella

_ obliquely cut off, so that one side of the umbrella is

longer than the other. Four radial canals. Four tentacle-
bulbs; three of which are without tentacles, and the other,
larger, with two tentacles situated on the longest side of
the umbrella. A medusa-bud is also present on the large
tentacle-bulb. The ex-umbrella has five longitudinal rows
of nematocysts, one following the course of each of the
three radial canals running to the non-tentacular bulbs»
and two near the canal which joins the large tentacle-bulb
carrying the two tentacles.

Steenstrup also found in the same locality specimens of
a small solitary hydroid, attached to the empty shells of

~Balanus, etc. The hydroid is about 4 inch long, with 5

or 6 tentacles in a single verticil upon the head. Below

the head four bell-shaped, quadrangular, meduse hang

down, at right angles to one another. The medusa has

the margin of the umbrella obliquely cut off. Four ten-

tacle-bulbs, without tentacles. The bulb on the longest
side of the umbrella is the largest. Steenstrup regarded
the large medusze with two tentacles as the adult stage
of the young meduse attached to the hydroid, and gave
the same name—viz., Coryne fritillaria to both of them.

Sars (20) found three specimens of a medusa, which he
named Steenstrupra globosa, in May, 1836, at Floroen in
Sondfjord. They resemble Coryne fritlaria in shape of
the umbrella, but have three tentacles on the large bulb
on the longest side of umbrella. In May, 1888, Sars
again found three more specimens; one with three
tentacles and the others with only one tentacle. Sars

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L.M.B.C. MEDUSA. et = OS

believed that these meduse were probably the same as
those taken by Steenstrup. The chief difference consists
in the number of tentacles. Heckel (75) considers that
they are distinct species and places them in his genus—
Amphicodon, which consists of three species—A. fritillaria,
A. globosus, A. amphipleurus.
. At Port Erin in 1893, on the first day of tow-netting,
(April 29th), I obtained specimens of Amphicodon. They
were fairly plentiful during the first few days of May and
disappeared about the 10th. Altogether I collected about
50 specimens. The umbrella is about 2 to 83 mm. in
length and not quite so broad. The opening of the
umbrella cavity is small as the velum is fairly wide, and
the large tentacle-bulb projects a little across the opening.
There are three non-tentacular bulbs and one large
bulb on the longest side of the umbrella which may carry
one, two, or three tentacles, usually two are present.
From the large tentacle-bulb meduse are budded off, and
often two, three or more buds are present in various
stages of development. I saw one young medusa break.
away from its parent, which it resembled in every detail
_ except size. The former was about 1 mm. in length
and the latter was nearly 8 mm. Both carried two
tentacles on the large bulb. The majority of specimens
were without medusa-buds and had ova or spermatozoa
upon the manubrium. ‘The sexes are separate. At the
end of March, 1894, I again visited Port Erin for the
purpose of collecting more specimens. The first specimen
was taken on March 29th, and during the whole of April
Amphicodon was very abundant in the Bay.

All the specimens have the five longitudinal rows of
nematocysts as described by Steenstrup. ‘The colour
of the tentacle-bulbs shows a considerable amount of
variation. In the young specimens yellowish brown is

—

ES SE ATT
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eS ES SR

ee a SE - ea
Se mS FSS ES a

| 254° TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the prevailing colour and in the adults a dark reddish -
brown. A few adult specimens were taken with brilliant
crimson tentacle-bulbs and with the ring and radial canals —
also full of a crimson pigment, not in circulation, but as —
small particles in the endoderm cells. These brilliantly
coloured specimens are an exception and duller colours
usually prevail. Another medusa—Huphysa aurata—
shows a similar briluancy of colour in a few specimens
only. | 3 | |

The most interesting feature connected with this’
medusa is the carrying of young hydroids in the umbrella
cavity. So far as I know this has not been recorded
before in any other medusa. I first observed this in the
specimens taken in 1893 which led me to obtain more
specimens in 1894 so that I might investigate the
development of the ovum. This I have nearly completed
by means of series of sections and the results will be
published soon. When the medusa first appeared at the
end of March the asexual method of budding medusz
from the base of the large tentacle alone existed, which
must add considerably to the original number of medusee
which leave the hydroid. Each medusa buds off at least

- four other meduse, all of which become sexual later on,

and some of the young medusz have also medusa-buds.
About the middle of April the reproductive cells began to
appear on the manubrium, at first as a slight swelling of a
transparent gelatinous appearance, faintly tinged with a
yellowish-brown colour. When squeezed, the mass breaks
up into small round cells (germinal cells), each with a
nucleus. As these cells increase in size a large oval mass —
is formed upon the manubrium. It appears from sections
that one of the original germinal cells becomes the ovum
which increases in size at the expense of the other germinal
cells which are absorbed. ‘The ovum remains attached to

2.M.B.C. MEDUSA. 255

the manubrium until the hydra-stage is reached. The
tentacles of the posterior verticil first make their appear-
ance about 11 to 17 in number, and when they become
about 2 mm. long, a second verticil of 8 tentacles appears.
The body of the young hydra behind the posterior tentacles
becomes covered with a thin perisarc. Sometimes the
young hydra breaks away from manubrium before the
appearance of the second row of tentacles and remains—
free in the umbrella cavity. The young hydra with the
two verticils of tentacles is about 2 mm. long, and the
tentacles of the posterior verticil are a little longer than
the body. Often in the same specimen there is an ovum
on the manubrium and two or three hydrae showing
various stages of development. In a few specimens the
asexual budding is taking place at the same time as the
sexual reproduction on the manubrium. Sometimes the
young medusa on the tentacle-bulb shows also the repro-
ductive cells on the manubrium. Several of the young
hydroids fixed themselves on the bottom of a glass
aquarium, and grew to about 10 mm. in length. A great
change took place in the body, which became stem-like and
the coenosarc showed longitudinal striae of an orange-
red colour. The eight oral tentacles increased slightly
in length and the whole head became larger. These
hydroids have every appearance of a young Corymorpha.
I can see no reason for specifically separating the free-
swimming medusae taken by Steenstrup off Iceland and
by Sars off Norway from the Port Erin specimens. It is
clear from the development of the hydroid of the Manx
meduse that the hydroid is not the same as Steenstrup
dredged off Iceland. It will be best to leave the Iceland
hydroid under the name of Diplura fritillaria Allman (4),
and regard Steenstrup’s free-swimming medusze with two
tentacles, and those taken by Sars with one and three
tentacles as Amphicodon fritillaria.

' 956 TRANSACTIONS LIVERPOOL, BIOLOGICAL SOCIETY.

-L. Agassiz (2) found in Massachusetts Bay a corymor-
pha-like hydroid budding off uni-tentacular medusz, to
which he gave the name Hybocodon prolifer. As Allman
(4) gives a description of the hydroid and medusa there is
no need to repeat all the details. The hydroid is about 2
inches long, about ~;th of an inch in diameter below the
head, and thinner near the base. Perisarc smooth except
near the head where there are a few annular constrictions.
Coenosarc of the stem with longitudinal orange-red striae.
Hydranth with two verticils of tentacles, about sixteen
tentacles in each. The hydroid lives in rock-pools on
out-lying rocks away from the shore, about the Laminarian:
Gone, usually three or four are together.

Between the two verticils of the tentacles the medusa-
buds are situated just asin Corymorpha nutans. Meduse
are budded off from January to April. Agassiz apparently
has not seen any specimens of the adult medusze but
only those either upon the hydroid or just liberated, and
his description is confined to the earliest stage of the
medusa. The umbrella of the young medusa has the
margin obliquely cut off so that one side of the umbrella
is longer than the other. On the longest side of the
umbrella is situated a large tentacle-bulb carrying a soli-.
tary tentacle, and also medusa-buds. There are three
other non-tentacular bulbs without tentacles in the usual
places upon the margin of the umbrella. The exumbrella
has five longitudinal rows of nematocysts which extend
from the margin to within a short distance of the apex of
the umbrella. One row is above each of the radial canals
running to the tentacle-bulbs not bearing tentacles, and
the other two rows are near the canal running to the
large tentacle-bulb. The rows of nematocysts are con-—
spicuous early in the year by their orange-red colour, but
become inconspicuous about April.

U.M.B.C. MEDUSA. Bene? oe ise

The young medusa Hybocodon prolvfer has all the charac-
teristic features of the medusa, Anyphicodon fritillaria.
The shape of the umbrella, the five rows of nematocysts,
and the budding of medusa from the base of the large
tentacle are common to both. The young hydroids
which I reared at Port Erin have every appearance of
becoming similar to the hydroid taken by Agassiz.
They agree in colour, in the longitudinal striae of the
stem formed by the coenosarc, and the tentacles of the
posterior verticil are about the same in number. The
anterior verticil has only eight tentacles, but others may
appear as the head grows larger. The discovery of the
adult hydroid with medusz on the Manx coast will settle
the identity at once.

. Bohm (Jena zeitschr. f. Naturw., Vol, XII, 1878)
captured a specimen of a medusa off Heligoland which
he believed to be identical with the American Hybocodon
prolifer. The medusa has two tentacles on the large
tentacle-bulb on the longest side of the umbrella and
many medusa-buds, and also the five rows of nematocysts
on the umbrella. Haeckel however, places Hybocodon
prolifer of Bohm as a synonym of Amphicodon fritullaria.

In the last Annual Report (1894) of the L.M.B.C., it
is stated on my authority that Amphicodon fritillaria is
an addition to the fauna of the British seas, but I have
recently read a paper by Mr. J. R. Green (13) on the
medusze on the coast of Dublin in which he clearly
describes specimens of Amphicodon fritillaria found on
that coast about 1856. He describes them as two new
species—Diplonema islandica and Steenstrupia owen.

Diplonema islandica, has two tentacles on the large
bulb on the longest side of the umbrella. Tentacle-bulbs
of a brilliant crimson colour. A medusa-bud on the large
tentacle-bulb, and in one specimen a medusa-bud on one

958 TRANSACTIONS LIVERPOOL. BIOLOGICAL SOCIETY.

of the tentacles, some distance from the base. Green
has read Steenstrup’s description of Coryne fritilaria
and states that there is a strong resemblance between the
two species, but does not think that they are the same.
Haeckel however believes that D.zslandica is the same
species as A. fritillaria. Steenstrupia owenw is very
much like D. islandica in shape. It differs from it in
possessing only one tentacle on the tentacle bulb. A
Stielcanal is also present, running from the top of manu-
brium to the ex-umbrella. Three medusa-buds at the base
of the large tentacle. Haeckel believes that this species
is probably the same as Hybocodon prolifer, Agassiz. It
is interesting to notice that these specimens were taken
on the coast of Ireland nearly opposite the Isle of Man.
Haeckel describes another species of Amphicodon—A.
amphipleurus, which he found on the coast of Normandy,
at Granville. It has three tentacles on the large tentacle
bulb on the longest side of the umbrella, and the usual
five rows of nematocysts on the ex-umbrella. It differs -
only from the other species of Amphzcodon in possessing a
larger number of medusa-buds on the large tentacle-bulb,
and in having a stiel-canal. The radial-canals, ring-canal
manubrium, tentacle-bulbs and tentacles of a crimson
colour. All the meduse of Amphicodon fritillaria must
have a Stielcanal on leaving the hydroid, but it usually
disappears later in life. I saw one adult specimen at Port
Hrin with a Stielcanal. In some species of meduse the
Stielcanal remains throughout life; in others 1t disappears,
but occasionally may be present in adult specimens.

Family TIARIDz.
Tiara pileata, (Forskal 15).
Oceania episcopalis, Forbes (9).
The medusa was first taken by Forskal in 1775.- Since

L.M.B.C. MEDUSA. 959 -

then it has been described under many names and by
many naturalists.

One specimen taken at Port Hrin, Aug. 1892. Fairly
abundant during May and June, 1893. Many specimens
were often left on the sandy beach by the tide. During
April, 1894, I obtained several young stages in the bay.
One specimen taken on April 18th, umbrella 7 mm. long,
and 4mm. wide. At the aboral end of the umbrella there
is a large globular mass of tissue, which looks like a knob
on the top of the umbrella. Four broad radial canals,
manubrium about one-third the length of the umbrella-
cavity, mouth wide with four large lips. On the margin
of the umbrella four perradial tentacles, of which two,
opposite ones, are larger and longer than the others. The
basal bulbs of these tentacles are large and long, each
with a reddish ocellus. There are also four interradial
tentacle-bulbs, with ocelli but without tentacles. <A
specimen taken on April 10th, with a short conical knob
at the aboral end of the umbrella. On the margin of the
umbrella there are four long perradial tentacles, four large
interradial tentacle-bulbs without tentacles, and eight
small adradial tentacle-bulbs. Another specimen taken
on April 10th, umbrella 4 mm. long and 23 mm. wide,
four perradial tentacles, 20 mm. long when fully expanded,
four short interradial tentacles, and eight adradial tenta-
cular-bulbs without tentacles. These three specimens
show clearly the increase of the tentacle-bulbs and
tentacles. In some specimens the number of tentacles
exceeds eighteen. Gosse records a specimen with twelve
tentacles and three tentacle-bulbs between each, the
central one being the largest. I have seen a specimen
with eighteen tentacles, and a tentacle bulb between every
two of them. Probably the maximum number is about
48 tentacles.

‘260 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Allman (Monogr. Gym. Hydr., p. 33, woodcut 8), figures
a young stage of Tiara under the name of Oceania coron-
ata. ‘his is probably a young 7. pileata. It has four
long, perradial tentacles ; four short interradial tentacles ;
and eight adradial tentacle-bulbs. It is exactly like a
specimen taken at Port Erin on April 10th. Heckel,
however, places this specimen under the name of Tvara
octona. i 7

Distribution — Mediterranean. Norway, Heligoland,
Shetland, St. Andrews.

Turris neglecta, Lesson (9). |

Byerley (6) records this species.—‘‘ Taken rarely in the
Mersey, by Mr. Price.”

I have not yet seen this species at Port Erin. It
appears to be a fairly common medusa and no doubt will
be recorded before long on the Manx coast. 3

Distribution.—St. Andrews, Southampton, Portland, —
Ilfracombe, Tenby, Liverpool.

Family MARGELIDA.

Dysmorphosa carnea, (M. Sars 15).

Podocoryne carnea, Hincks (16), Allman (4). :
One specimen taken on March 30th, 1894, with four
long perradial and four short interradial tentacles.

Distribution.—Naples, Norway, St. Andrews, Firth of
Forth, Plymouth, Falmouth.

Dysmorphosa muuma, Heckel (15).
This medusa was first discovered by Heckel in 1865
off Heligoland, and has not been recorded since. The
length of the umbrella is 0°8 mm. and the width 0°6 mm.
Four simple oral tentacles terminating with a cluster of
nematocysts. Medusa-buds upon the manubrium. Hight —

L.M.B.C. MEDUSZ. 261

tentacles upon the margin of the umbrella, 4 single
perradial and 4 single interradial. Blackish tentacle-bulbs.
Heckel’s figure of Dysmorphosa nunivma shows a con-
striction across the umbrella-cavity. Ido not think this
is a permanent constriction, but produced probably by
injury or death. In all medusz.which have a permanent
constriction across the umbrella, the constriction is either
level with the top of the umbrella-cavity or above it, so as
not to interfere with the size of cavity and the contractions
of umbrella. I have seen meduse under the influence of
cocaine die with a constriction across the middle of the
umbrella-cavity. In the museum of the Marine Biological
Association at Plymouth I saw specimens of Dysmorphosa
minima, but labelled Dysmorphosa carnea. Nine of the
Plymouth specimens correspond to Heckel’s description
and five others only differ in the number of tentacles.
These show an increase in the perradial groups and have
the tentacles arranged thus :—
Perradial 2211 | 2121 | 1112 | 2229
itapemradial 1111 | W111 | bit) 1111

The perradial groups with two tentacles have one
tentacle longer than the other, the smaller one 1s evidently
srowing. I took a single specimen at Port Hrin, 1894,
of a medusa which is exactly lke the specimens of
Dysmorphosa mimma at Plymouth. It has four simple
oral tentacles; eight single marginal tentacles; and four
medusa-buds upon the manubrium.

It is quite possible that Dysmorphosa minima is an
early stage of Lizzia blondina. 'The Plymouth specimens
clearly show an increase of tentacles in the perradial
groups.

Dysmorphosa nunima may easily be distinguished from
Dysmorphosa carnea by the stomach being attached to a
short peduncle, and also by the presence of medusa-buds
upon the manubrium,

962 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

2 Cyteandra areolata, (Alder 15).
Hydractinia areolata, Alder (8).
~ Podocoryne areolata, Hincks (16), Allman (4).

1 ‘site temporarily under the name of Cytwandra
-areolata, until further evidence can be obtained, a number
of meduse taken at Plymouth and Port Erin. They all
belong to the same species and show different stages of
development. The youngest stage 1s somewhat lke
Cyteandra areolata. The uncertainty mainly arises on
account of the great rareness of the hydroid stage and the
plentifulness of the meduse in Port Erin Bay, Alder first
described the hydroid stage under the name of Hydrac-
tuma areolata. He obtained his specimens on a dead
shell of Natica aldert, brought in by a fishing boat at
- Cullercoats, and Norman sent him another specimen from
-Shetland. Alder has seen the young meduse leave the
colony. The medusa has a moderately deep, subglobose
umbrella, with nematocysts scattered upon the outside;
four radial canals; four short perradial and four shorter
interradial tentacles, and eight adradial bulbs on the
margin of the umbrella; four clusters of nematocysts
near the mouth. es

Alder is the only person who has seen the young med
-and since his observations no one has recorded this
species. Huincks states that Hodge has taken the hydroid
“with meduse at Seaham. Allman, however, omits this
reference in his monograph, and I cannot find any state-
‘ment referring to this species in Hodge’s papers.

Hincks removed Hydractinia areolata to the genus
Podocoryne, where it still remains. There are three
species of Podocoryne recorded for Great Britain—P.
carnea, P. proboscidea and P. areolata. Podocoryne carnea
is the commonest; P. proboscidea was taken ee Hincks
-at Ilfracombe, but has not been since recorded.

L.M.B.C. MEDUSA. 263

IT had at Plymouth a colony of P. carnea in a bottle.
It budded off many meduse which were not like the figure
of the medusa of Podocoryne areolata given by Alder, nor
like the meduse which I have taken at Port Erin. The
Port Erin specimens approach nearer to Alder’s figure of
P. areolata than to any other known medusa. I prefer
to leave them for the present under this name rather than
to make a new species.

The earliest stage of the series I captured in Port etn
Bay on March 31st, 1894. The umbrella about as long
as broad, 1} mm. in diameter; the ex-umbrella covered
with nematocysts. There are sixteen tentacles—4 perra-
dial, 4 interradial and 8 adradial. The perradial tentacles
are longer than the interradial, and the latter are longer
than the adradial. The large bulbs at the base of the
tentacles, contain a reddish-yellow pigment. In addition
to the tentacles there is one small bulb placed between
two tentacles, which is evidently the commencement
of another tentacle. Near the mouth there are four
simple oral tentacles, each terminating in a cluster of
nematocysts.

This naturally appears to be a later stage os Alder’s
medusa of P. areolata. The main difference consists in
the presence of adradial tentacles in the place of the bulbs.
It is, however, usual for tentacles first to appear as small
bulbs upon the margin of the umbrella. The next stage
was taken at Plymouth, September 30th, 1893. The
umbrella nearly as long as broad, about 1 mm. in diameter,
the ex-umbrella covered with nematocysts. On the
margin of the umbrella there are 16 large tentacles—4
perradial, 4 interradial and 8 adradial. The tentacle-bulbs
at the base of the tentacles of a dark brown colour. In
addition to the large tentacles there are four small, slender
tentacles, with colourless tentacle-bulbs, These are

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264 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

. evidently in a state of development. Four simple oral
_tentacles with clusters of nematocysts.

A specimen taken at Port Erin, April 18th, 1894.
Umbrella about 3mm. in diameter. Sixteen large ten-
tacles and six small ones. The oral tentacles are once
dichotomously divided, each branch terminating in a
cluster of nematocysts. It has been observed in several
species of medusze which have branched oral tentacles in

the adult stages that they start life with simple, unbranched

tentacles. In Margelis the number of branches increases
with age.
Two specimens taken at <Pawt ast June 5th, 1893,

about 2mm. in diameter. Sixteen large tentacles, with

reddish brown tentacle bulbs, and ten small tentacles,
with faintly coloured brownish tentacle-bulbs. Four oral
tentacles, once dichotomously divided, with terminal
clusters of nematocysts.

A specimen taken at Plymouth, Septerites 25th, 1893.

Umbrella about 1 mm. in diameter. Sixteen large
_tentacles with dark brown tentacle-bulbs, and ten small

tentacles. Hour unbranched oral tentacles. .
Three specimens taken at Port Erin, April 27th, 1804.
Umbrella about 4 mm. in diameter, with 25, 28, and

. 80 uniform tentacles. Tentacle-bulbs dark red and very

conspicuous. Four oral tentacles once dichotomously
divided. Other specimens taken on May 5th and 9th
somewhat similar to the above.

The youngest medusa of this series starts with sixteen —

tentacles and a small bulb. The other specimens show a

gradual increase of 20, 22, 25, 26, 28 and 30 tentacles, the

maximum number is probably 32. The young tentacles

increase in size. In some of the largest specimens all

_the tentacles are alike in colour and size. .The~ oral

at first have colourless bulbs, but colour appears as they —

L.M.B.C. MEDUS&. 265
tentacles at first are unbranched and become branched
later in life. The umbrella in all specimens is about as
long as broad, in the older specimens the aboral end has
a thick mass of tissue. The ex-umbrella is covered with
nematocysts. This medusa may easily be identified, as
it is the only British medusa with oral tentacles which
has sixteen or more tentacles fairly evenly distributed on
the margin of the umbrella. The other meduse with
oral tentacles have either less than sixteen tentacles, or a
large number which are grouped into either four or eight
bundles. 7

Inzzia blondina, Forbes (9).

Forbes first described this species from specimens taken
in the Sound of Brassay, Shetland, 1845.

I took some specimens in Port Erin Bay at the
beginning of May, 1894.

It is difficult at first to distinguish this species from
the early stages of Margelluwm octopunctatum.

The umbrella is somewhat like Margelliwm. octopuncta-
tum in shape but smaller in size, usually about 13 mm.
long and 1} mm. wide. On the margin of the umbrella,
in the adult stage, there are three tentacles on each of
the four perradial bulbs and only one tentacle on each of
the interradial bulbs. The single interradial tentacle is one
of the characteristic features of this species. In Margel-
lium octopunctatum usually two or three tentacles are
present on the interradial bulb. Lizzia blondina has
never more than one interradial tentacle on each bulb.

The tentacle-bulbs have a hght yellowish colour, but
the tentacles are colourless.

There are four simple oral tentacles, each terminating
in a single cluster of nematocysts. ‘These never have
two terminal clusters as in Margelliwm octopwnctatum.

966° TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Margellium begins to appear early in the spring, usually
abundant in April, and becomes scarce during May.
Lizzia begins to appear about: Mey Forbes states that
it is of solitary habit.

Ihave taken specimens with two tentacles in each of |
the perradial groups. Medusa-buds are usually present
on manubrium, at first yellowish in colour.

It is possible that Dysmorphosa minima is the earliest
stage of this medusa.

Distribution.—Shetland, St. Andrews, Cornwall.

Margelis principis, Steenstrup (15).

This medusa is not nearly so abundant as Margelis
ramosa. It may be easily recognised and distinguished
from M. ramosa by the very large crescent or U-shaped
tentacle-bulbs. It also has about twice as many marginal —
tentacles in each group and about three times as many
branches on the oral tentacles. Steenstrup first described
this medusa’ from specimens obtained off the Faroe Islands
and Iceland. Heeckel describes and figures the original
specimens taken by Steenstrup which are in the museum
at Copenhagen.

- In 1893 I met with a single specimen of this species. It
was taken in Fleshwick Bay, near Port Erin, on May 21st.

The Manx specimen agrees with Heckel’s description
except in some minor details. Heckel figures the tentacle-
bulbs with a dark -horizontal line at the top and under it
an ornamental wavy line. The Manx specimen has in
place of the wavy line a series of longitudinal lines, one
opposite each tentacle. At the base of each tentacle there
is a black ocellus. About 25 tentacles in each group and
about 25 terminal branches to each oral tentacle. Um-
brella 4 mm. in length and width. On May oth, 1894,
I met with another specimen in Port Erin Bay. The

L.M.B.C. MEDUSE. Oey

umbrella 4 mm. in diameter. About 15 tentacles in each
group, the outside ones being very small and evidently
erowing.

The compound tentacle-bulbs are curved, but not so
much as in the adult. The upper portion of the bulb
contains a dark brown pigment and from this mass
longitudinal bands extend towards each tentacle. A black
ocellus at the base of each tentacle. The oral tentacles
with about 16 branches. Generative products not present.
This is evidently a young specimen. It may be difficult
to distinguish the early stages of M. principis from those
of M. ramosa. There is a specimen of M. principis in the —
museum of the Mar. Biol. Assoc. at Plymouth. Romanes
(18) has taken specimens in Cromarty Firth (= Bou-
gamnvillea. allman et fruticosa, Romanes).

Margelis britanmca, (Forbes 9).
Bougainvillea britanmca, Forbes (9).
Margelis ramosa, Heckel (15).

Forbes first discovered and described this medusa in
1841, from specimens taken off Bute. Since then it has
been recorded from many places along our coasts. Clubb —
(7) found specimens in Port Erin Bay during August, 1885.

The umbrella is about as long as wide, 4 to 6 mm.
Four perradial groups of tentacles with 8 to 16 tentacles
in each group. The tentacles are united at the base and
form a compound tentacle-bulb. Near the base of each
tentacle there is a conspicuous ocellus. The oral tentacles
are usually three times dichotomously divided, and each
of the eight terminal branches ends in a cluster of
’ nematocysts.

During May, 1893, I collected many specimens in Port
Erin Bay. The compound tentacle-bulbs are yellowish,
and the tentacles, when expanded, about twice the diameter

268 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of the umbrella. A specimen taken on June 5th was
beyond the average size. It measured 8 mm. in length
and 6; mm.in width. About 12 marginal tentacles in
each of the four groups. The oral tentacles five times
dichotomously divided. During March and April, 1894,
I took with a tow-net some early stages of this medusa.
The earliest stage was taken on March 29th. The um-
brella about 1 mm. in diameter, with a broad velum.
Four marginal groups of tentacles; two of which have
three tentacles and the others have only two. At the
base of each tentacle there is a crimson ocellus. The
compound tentacle-bulbs with two tentacles have also
three ocelli. The extra ocellus belongs to the third ten-
tacle which has not yet developed. The oral tentacles
also show an early stage of development, only three
branches, each terminating with a cluster of nematocysts.
Another specimen shows a further development of the
marginal and oral tentacles. There are three tentacles
‘in each of the marginal groups, but they are unequal in
length, one of the outside ones being about half the length
of the other two. At the base of each tentacle there is a
red ocellus. The compound tentacle-bulbs yellowish
brown. ‘The oral tentacles have four branches.

A specimen taken on March 31st has the central tenta-
cle of the three marginal tentacles the shortest, which
shows that the third tentacle in each group may either
develop on the outside of the two primary tentacles, or
between them.

A specimen taken on April 3rd, about 2 mm. in diame-
ter, with five tentacles in each marginal group. The three
central ones large and about equal in length, and two, one
on each side of the central three, just beginning to develop.
The oral tentacles have five terminal branches, each ending

in a cluster of nematocysts. On May 9th a specimen

L.M.B.C. MEDUSA. 269

was taken, about 4 mm. in diameter, with eight tentacles
in each group. Black ocellus at the base of each tentacle.
The oral tentacles have eight terminal branches, each
tentacle being three times dichotomously divided.

These specimens illustrate very well the increase in
number both of the marginal tentacles and of the branches
of the oral tentacles. The young medusa usually leaves
the hydroid with two tentacles in each group, and the
oral tentacles are either simple (unbranched) or with a
single branch. |

On April 28th I captured two specimens, which are
certainly an early stage of a Margelis. The umbrella
about 3 mm. in diameter. _The umbrella cavity is almost
_ obliterated by the thickness of the umbrella. Three
marginal tentacles, unequal in size, in each of the four
groups. One specimen has one oral tentacle unbranched,
and the other three with one branch. I think these are
probably the young stages of Margelis ramosa, with an
abnormal growth of the umbrella.

The medusa Bougainvillea britannica, Forbes, is the
adult stage of the medusa which is liberated from the
_hydroid Bougainvillia ramosa, Van Beneden. Wright
(22) in 1857 succeeded in rearing meduse from the hydroid
colonies of Atractylis ramosa (=Bougainvillea ramosa,
Van Beneden). The young meduse first had four simple
oral tentacles, and eight marginal tentacles, two in each
group. Stomach and tentacle-bulbs of an orange colour.
A black ocellus at the base of each tentacle. Wright was
able to keep his specimens alive and saw further develop-
ment, The oral tentacles became first once, and after-
wards, twice dichotomously divided. Additional tentacles
also developed in the marginal groups. The greatest
number observed was six, and each one had a black ocellus
at its base. At the same time the generative products

270 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

began to develop on the stomach and also to extend for a
short distance along the radial canals. Wright obtained
young medusze from his colonies in the spring and
‘autumn.

-Distribution.—Belgium, Heligoland, Shetland, ie.
marty Firth, St. Andrews, Firth of Forth, Pr
Bute, Ballycastle Bay, Dalkey, Kingstown.

Margelliwm octopunctatum, (Sars 15). ~
Cytais octopunctata, Sars (19).
. Lnzzia octopunctata, Forbes (9).
Rathkea octopunctata, Heckel (15).

Sars first. discovered this medusa on the Norwegian
coast, and described the species with three tentacles in
each of the perradial and interradial groups, and. with
medusa-buds upon the manubrium. The young meduseze
on leaving their parent have 3 tentacles in each perradial
eroup and four single interradial tentacles.

- Forbes first added the species to the list of British
Meduse and captured the specimens off the Shetland —
Isles. He observed that all his specimens had three
tentacles in each perradial group and either two or three
tentacles in the interradial groups. Forbes considered —
his specimens to belong to the same species as those
described by Sars. Heckel has divided the specimens
taken by Sars and Forbes into two different genera, on
account of some of Forbes’s specimens having only two
tentacles in each interradial group. Meduse with an equal
number of tentacles in the perradial and interradial groups
belong to the genus Rathkea and under R. octopunctata
Heckel places Cytwis octopunctata, Sars. .Meduse with
more perradial than interradial tentacles belong to the
genus Margelliwm, and as some of Forbes’s specimens
have only two tentacles in the interradial groups Heckel

ECD CL MEDUS mye s 5 ipo ee TEL
has removed Lizzia octopunctata, Forbes, to the genus
Margellium.

During May, 1893, I found in Port Erin Bay medusz
which belong undoubtedly to the same species as those
described by Forbes as Lizz7a octopunctata. Some of the
specimens correspond exactly with the descriptions given
by Sars and Forbes, and others are without doubt other
stages, showing earlier and later forms, of the same
species. The oral tentacles of all the specimens taken
have four clusters of nematocysts. Each oral tentacle
bifurcates near its extremity and each branch terminates
with a cluster of nematocysts. The two other clusters
are on a short branch, one on each side of the main stem
of the tentacle. | lay te

The umbrella of the smaller specimens is from 1 to 2
mm. in diameter, and of the larger specimens about 3 to 3}
mm. in length and slightly less in width. The colour
of the tentacle-bulbs varies, being either yellowish brown,
dark brown, or black. The great difference in the speci-
mens is in the number of tentacles in each group, and it
is upon the number of tentacles that Heckel mainly bases
his” classification. Twenty specimens are sufficient to
show the change of number of the tentacles in each
group, and in the following list I have arranged 20 speci-
- mens so as to show the increase in the number of tentacles
in each group. This increase is not a variation, but
_sunply shows stages in development.

Number of tentacles in
Number of specomens. | each group.
1. Perradial—3333 Margelluum stage (Heckel).
Interradial—2222 Liazva stage (Horbes).
4. Perradial—3333 | 3333 | 3333 | 3333
Interradial—a3332 | 3233 | 3233 | 3233

272 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

3. Perradial—3333 Rathkea stage (Heckel).
Interradial—3333 Inzzva stage (Forbes).
5. Perradial—3334 | 4333 | 4838 | 3484 | 3444
Interradial—3333 | 3333 | 3833 | 3333 | 3333
2. Perradial--4444
Interradial—3333
4, Perradial—5443 | 5444 | 4354 | 4554
Interradial—3333 | 3333 | 3333 | 3333
1. Perradial—5555 ? Margellium gratum,
Interradial—3333 | Heckel.
Sars states that the young medusa leaves its parent
with 1 interradial and 3 perradial tentacles. This stage I
did not meet with, but found the next stage with 2 inter-
radial tentacles. ‘The interradial tentacles never exceed

three in each group, but the perradial tentacles increase
until five is reached, which is the maximum. Agassiz (J)
describes an American species Lizzia grata, (Margelliwm
gratum, Heeckel), which has five perradial and three
' interradial tentacles in each group. ‘These tentacles
develop in the following order. At first there is in each
perradial group one long median tentacle, and on each
side of it a small tentacle. Next, the median tentacle of
the interradial group appears, followed by a pair, one on
each side of it. At this stage the medusa has 3 tentacles
in all the groups. Finally the two remaining tentacles of
the perradial groups appear to complete the set of five.
Agassiz also describes the development of the oral ten-
tacles. In the young medusa there are four simple,
unbranched, oral tentacles terminating in a cluster of
nematocysts. Each tentacle branches near the extremity
and each branch terminates with a cluster of nematocysts.
Next the lateral clusters appear, one on each side of the
original tentacle. Then arises a cluster midway between
the bases of the four tentacles, and finally, a cluster
between the branches at the end of the tentacle,

L.M.B.C. MEDUSA. 273

The Port Erin specimens with 5 perradial and 3 inter-
radial tentacles in each group, are very much like the
American species except that the oral tentacles have fewer
groups of nematocysts. The other groups of nematocysts
may have yet to appear, if so, there will be no reason for
separating the English and American species. It is best
however, for the present, to keep them apart until more
is known about the English species. On my arrival at
the Plymouth Laboratory in September, 1893, I found
that Mr. Garstang had also been collecting specimens of
Lizzva octopunctata, Forbes, during March and April in
the Sound. He believes that Rathkea octopunctata and
Margelliwm octopunctatum are the same species, and has
recently published his notes (12) which show the develop-
ment of the marginal tentacles. In the museum of the
Marine Biological Association at Plymouth there are
two dozen specimens of Margellium octopunctatum col-
lected by Mr. Garstang, all the specimens, except two,
have 3 tentacles in all the groups. The two exceptions
have their tentacles arranged thus:

iPerradial soc |. o222:
apermadial Joo | alli,

As none of Garstang’s specimens have more than three
tentacles in each group, he regards M. octopunctatum as
an early stage of Rathkea octopunctata, and the former
name is done away with. But the specimens taken at
Port Erin show that a further increase does take place in
the perradial tentacles, and that there are more perradial
tentacles than interradial, which is the characteristic
feature of the genus Margelliwm. Therefore I retain the
name of the genus Margelliwm.

On Feb. 24th, 1894, I received from Mr. Garstang
seven specimens, alive, of the early stages of Margelliwm

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_ 274 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

_octopunctatum. The number of tentacles in each BrOuy
is as follows:
3 specimens—Perradial 3338.
Interradial 1111. =e
The oral tentacles are without the lateral clusters of

nematocysts, but one specimen has a few nematocysts

which mark the position and origin of these clusters :
3 specimens—Perradial 3333 | 3333 | 3333.
seek Interradial 1112 | 1312 | 2282.
The second tentacle in the interradial groups is very

small compared with the other, and has. just commenced
to grow.
All the specimens have medusa-buds upon the manu-
brium. Tentacle-bulbs yellowish brown. Diameter of the
umbrella about one millimetre. 3 7
At Port Erin on the 22nd March, 1894, the first ae of ;

tow-netting, I found early stages of Margelliuwm octopunc-

tatwm, and on every occasion on which the townet was
used specimens of this medusa were taken. At the end
of April it was very abundant and the specimens had |
reached the Rathkea-stage, with three tentacles in all the
groups. As I left Port Erin at the beginning of May I
did not meet with the later stages which I obtained at
the end of May and the beginning of June in the previous
year. Mr. Garstang obtained his specimens at Plymouth
during February and March, and the medusa disappeared

in April, which probably accounts for his not obtaining

specimens with more than three tentacles in each group.
T took many specimens similar to those taken in the
previous year, showing a similar arrangement in the
eroups of tentacles. There is no need to repeat the list
again. One specimen however, is exactly like one taken |
at Plymouth on Feb. 24th, with the tentacles arranged
thus—perradial 3333, interradial 2111. I think that the

L.M.B.C. MEDUSA. Q75

Plymouth and Port Erin specimens belong to the same
species.

_ The budding of medusze from the manubrium is not the
sole means of reproduction. The sexual method comes
on later in life, and the sexes are separated. The asexual
method of budding takes place first and is followed by
sexual reproduction.

The same thing occurs in Amphicodon fritillaria, which
at first buds off meduse from the base of the tentacles
and then usually ceases to reproduce asexually as soon as
the sexual cells begin to appear upon the manubrium.
It is quite possible that some of the Sarsidze which are
now regarded as distinct species on account of their
budding off meduse may only be stages of other species
which are described as sexual.

An abnormal specimen of this species was taken in Port
Hrin Bay on May 10th, 1893. It has six radial canals

instead of the normal four. The additional canals run

down to two adjacent interradial groups of tentacles,
which possess an abnormal number of tentacles, four
being present, while in normal specimens the number
never exceeds three. The number of tentacles in each
group is as follows :—Perradial 3358, Interradial 2441.
There are five oral tentacles. Four of these have four
clusters of nematocysts, two terminal and two lateral.

The other has a single terminal cluster and two lateral

clusters. An abnormal number of radial canals is very
rare among the Hydromeduse.

Distribution.—Norway, Shetland, St. Andrews, Fal-
mouth, Plymouth.

Thaumantias hemisphaerica, (Muller).
Thaumantias punctata, Forbes (9).

_ Clubb reports (7) that 7. hemisphaerica was abundant

276 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

in Port Erin Bay in August, 1885. This species has been
recorded in nearly every list of British meduse. Up to
this time I have not met with a single living specimen of —
this common medusa. Every specimen taken, which
has any resemblance to description and figure of this
species, possesses marginal vesicles between the tentacles.
Heckel has placed T. hemisphaerica, Muller, among the
Thaumantide, a family which is characterised by the
absence of marginal vesicles between the tentacles.

I may here say that specimens preserved in spirit are
not to be relied upon for the absence of marginal vesicles.
The otolith usually disappears and the vesicle shrivels up.

Forbes obtained specimens of TJ. punciata off the Isle

of Man in June, 1839. Heckel considers this species to
be a form of T. hemisphaerica. }

Laodice cruciata, L. Agassiz (15).
Thaumantias pilosella, Forbes (9). |

Garner (10) found this species abundantly in Douglas
Bay.

Laodice cruciata is recorded by me in the Severn
Annual Report (1898) of the L.M.B.C. amongst the species
taken at Port Erin. I regret to say this 1s an error on
my part, as: the specimens have since been identified as
Mitrocomella polydiadema.

This species is widely distributed, Naples and oie
places in the Mediterranean, Holland, Shetland, St.
Andrews, Plymouth, Falmouth.

Laodice calearata, Li. Agassiz (2).
Lafoea calcarata, (A. Agassiz 1).
On May 5th, 1894, I took in Port Erin Bay a single
specimen of a medusa which I believe to be an early stage
of Laodice calcarata, an American medusa which so far

L.M.B.C. MEDUSA. DLT

as I know has not been recorded for Europe. A. Agassiz
gives an excellent description of the life-history of this
species and also a description of its hydroid.

Umbrella flatly-arched, about twice as broad as long,
stomach flat; mouth with crenate lips. Four radial canals.
The gonads extend nearly along the whole length of the
canals, starting from stomach and reaching nearly to the
marein, forming in the adult specimens a folded band.
On the margin of the umbrella 100 to 200 tentacles.
Between these tentacles there are short filiform cirri, and
also club-shaped cirri. No marginal vesicles. In the
- basal bulb of the large tentacles there is a black ocellus.
The largest specimens are about an inch in diameter.
Umbrella and tentacles colourless. Ovaries yellowish.
The Port Erin specimen is only 5 mm. in diameter, with
about 30 tentacles, and cirri of both shapes. It corresponds
to the description given by Agassiz. It is certainly not
Laodice cruciata.

Agassiz obtained his specimens in Buzzard Bay, North
America, 1861.

Melicertidiwm octocostatum, (Sars 15).
Stomobrachwum octocostatwm, Forbes (9).

Two specimens taken at Port Erin on May 27th, and
one on June 2nd, 1893. Forbes describes the tentacles as
being uniform, but Ehrenberg and Romanes (19) have
seen a small tentacle between two large ones. The Port
Erin specimens have also one small tentacle, sometimes
two, between every pair of large tentacles.

This species may easily be identified by the eight radial
canals. :

Distribution.—Norway, Cromarty Firth, Ireland (N.
W. Coast), Bute, Arran, St. Andrews.

{
{
!

278 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.
Family Eucopipé.

Clytria gohnstont, Alder (16).

Several colonies of the hydroid stage were anegeee off
Port Erin on March 24th, 1894. They were placed in a
vessel of sea-water, and for many days medusze were
budded off in large numbers. The young medusa corres-
ponds exactly to the description and figure given by
Hincks. I tried to rear the medusz, but failed to get
them to live beyond a few days. I also obtained a few
specimens with the tow-net in the bay at the end of
March. One specimen taken on March 25th had a single
adradial tentacle-bulb, and the interradial tentacle-bulbs
showed signs of budding out tentacles.

Hucope octona, (Forbes 15).
~Thaumantias octona, Forbes (9).
Clubb (7) records this species as fairly common off Port
Erin, August, 1885. ES Bee”
Distribution.—St. Andrews, Cornwall, Oban, Arran,
Tarbert (Loch Fyne). |

Obelia luctfera, (Forbes 15).
Thaumantias lucifera, Forbes (9).

I have taken specimens in Port Erin Bay during August,
1892; May, 1893; and April, 1894. Professor Herdman
obtained specimens off Port Erin in August, 1885. ;

, Distribution.—Heligoland, Shetland, St. Andrews, Dart-

mouth, Plymouth, Lizard Point, Hebrides. “

Tiaropsis multicirrata, (Sars 15).

- Thaumantias melanops, Forbes (9). 4
Three specimens taken in Port Erin Bay on May 6th,
1893. The largest 7 mm. in diameter, with about 180 :
tentacles. 8 marginal vesicles, with a black ocellus atthe
base of each,

L.M.B.C. MEDUSA. 279

At the end of March, 1894, I took with the tow-net
some early stages. One specimen taken on March 30th.
Umbrella bell-shaped, about $mm. long. 24 tentacles—
6 in each segment. 8 adradial marginal vesicles, each
containing 3 otoliths. Tentacle-bulbs of a brownish
colour. Another specimen taken on March 31st about 30
tentacles present. A black ocellus at the base of each
marginal vesicle. A specimen taken on April 18th with
80 tentacles. - |

Distribution.— Norway, MHeligoland, Shetland, St.
Andrews, Cromarty Firth.

' Hpenthesis cymbaloidea, (Slabber 15).
Thaumantias thompsont, Forbes (9). .
Clubb (7) records that this species was obtained in
fairly large numbers, during August, 1885, off Port Erin.
. Distribution —France, Holland, Cornwall, Roundstone
(Ireland).

Mitrocomella polydiadema, (Romanes 15).
Tiaropsis polydiademata, Romanes (18).

Romanes first gave a correct description of this medusa
from specimens taken in Cromarty Firth in 1875; and it
has not been since recorded. Romanes describes the
species as follows:—The umbrella resembles a deeply
shaped bowl. Manubrium so small as to be almost
invisible. Tentacles 45 in number and arranged in two
series, in one of which the tentacles are long and in the
other short. Four radial canals. Sixteen marginal
vesicles, 4 in each quadrant, and each contains about 30
otoliths. The medusa is brilliantly luminous when stim-
ulated, the hight being confined to a narrow and continuous
line round the margin of the umbrella. The manubrium,
radial canals, ovaries and tentacles of a rich rose colour,
Umbrella about 1 inch in diameter,

980 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY,

During May, 1893, I captured many specimens in Port
Erin Bay. On May 30th many dozens were left by the
tide upon the sandy beach, and amongst the weeds in the
rock pools. The largest specimens were about 30 mm. in
diameter.

At the end of April, 1894, I obtained specimens of some
early stages. ‘I'he smallest measured 4 mm. long and 43
mm.wide. Umbrella bell-shaped. Sixteen large tentacles,
4 in each quadrant, and between every two of them either
3 or 4 smaller tentacles. Sixteen marginal vesicles, some
with otoliths arranged in two rows, others with scattered
otoliths. The gonads just commencing to develop along
the radial canals. When the gonads are ripe, the genital _
band is folded.

Another specimen 12 mm. wide and 8 mm. long.
Umbrella globular. Manubrium very short; mouth with
four lips. The genital band folded along the radial canals.
Sixteen marginal vesicles, each with 15—20 otoliths
arranged in two rows, the largest being in the centre.
About 35 large tentacles and from 3 to 6 smaller tentacles
between every two large ones. The large tentacles when
fully expanded are about twice the length of the umbrella.
Mouth and tentacle-bulbs of a purplish colour. [2
bands yellowish brown, sometimes purple.

This medusa may be distinguished by the presence of
16 marginal vesicles, which are usually hidden by the
small tentacles.

Phialidium variabile, Heckel (15).

At the beginning of May, 1894, I captured two specimens
of a medusa which I believe to be Phiahdium variabile.
The first taken on May 5th. Umbrella bell-shaped, 10
mm. long and 13 mm. wide. 29 tentacles, eight in three
of the quadrants and five in the other. Two or three mar-

we a le ae ial alll

ae eT ee
'

L.M.B.C. MEDUSA. O81

ginal vesicles between every two tentacles, with a single
otolith in each. Stomach very short, and mouth with
four lips. Reproductive organs occupying the lower half
of the four radial canals, and slightly folded into a longi-
tudinal wavy line. Tentacle-bulbs and reproductive
organs yellowish brown.

The second specimen taken on May 9th. Umbrella if
mm. long and 21 mm. wide. 388 tentacles, ten in three
of the quadrants, and eight in the other. Two or three
marginal vesicles between every two tentacles with a
single otolith in each. Reproductive organs occupying
the lower half of the radial canals. Tentacle-bulbs and
reproductive organs yellowish brown.

On May 19th, 1893, I took a medusa in Port Erin Bay
which I also believe to be Phialidiwm variabile. Um-
brella about 9 mm. long and 12 mm. wide. 32 tentacles,
nine in two of the quadrants, and seven in the other two,
a single marginal vesicle between every two tentacles,
with a single otolith in each. Reproductive organs
occupying the lower half of the four radial canals, long
and oval in shape, and of pale greenish colour. 'Tentacle-
bulbs yellowish. This medusa corresponds with Forbes’s
description and figure of Thawmantias inconspicua which
appears to be a somewhat earlier stage, about 18 mm. in
width, 16—20 tentacles and between each pair ‘‘a rudi-
mentary marginal tubercle’ (marginal vesicle). Ovaries
long and linear, and of a faint lilac or greenish hue,
occupying more than half the length of each radial canal.
Tentacle-bulbs yellowish. Heckel, however, regards
Thaumantias inconspicua, Forbes, as a stage of Thauman-
tias henispherica, Eschscholtz (= T. hemispherica,
Forbes). Heckel has placed 7. hemispherica amongst
the Thawmantide a family distinguished by the absence
of marginal vesicles, M‘Intosh (17) appears to regard

282 TRANSACTIONS. LIVERPOOL BIOLOGICAL SOCIETY.

T. inconspicua, Forbes, as a variety of Phialidiwm varia-
bile, which was abundant at St. Andrews in June, 1889.
None exceeded a quarter of an inch in diameter, and all
were immature.

Clubb (7) records Thawmantias convexa, Forbes (9)
which was found by Mr. I. C. Thompson, off Penmaenmawr, ~
in July, 1885. Heckel considers 7’. convexa-to be identical
with Phialidium variabile.

Hutuma msignis, (Keferstein 15).

Keferstein first described this medusa from St. Vas
Normandy, 1862. In 1893 I took an adult specimen at
Plymouth. It is a rare medusa and has not been
recorded from other localities. |

In the adult the umbrella is hemispherical, about 7 mm.
wide and 3 mm. high. The manubrium is very long,
about 9mm. Mouth with four large lips. The gonads
are on the radial canals, along the lower half of peduncle.
On the margin of umbrella there are four very long
perradial tentacles, about 20 mm. long when fully ex-
panded. At the base of each tentacle there is a pair of
cirri. About 80 bulbs are also evenly distributed upon the ~
margin, each with a pair of cirri. Hight adradial mar-
ginal vesicles, each with 2—5 otoliths. The medusa is
perfectly colourless. |

An early stage of this medusa was taken in Port Hrin
Bay, 1893. Umbrella about 13 mm. in diameter. Four
large perradial tentacles, four interradial and eight adra-
dial bulbs. A pair of cirri at the base of each tentacle and
bulb.

Saphenia manos, (Wright 15).
First taken by Strethill Wright in 1858 near Queens
ferry, Firth of Forth.

an
‘ 4 y

L.M.B.C. MEDUSA. 283.

The umbrella of an adult specimen is about one inch
in diameter. Two large, colourless tentacles on the
margin of the umbrella, when fully expanded about 24
inches long, and also about 100 small bulbs each with
two minute cirri, armed with nematocysts. Hight mar-
ginal vesicles, each with four otoliths. The stomach is
on a peduncle about 1$ inches long. Four radial canals.
At Port Erin in 1893, I captured 3 specimens of the early
stages. One specimen about 1mm. in diameter. Two
long opposite perradial tentacles. Two perradial, four
interradial, and four adradial bulbs, all with a pair of
cirri. The two other specimens slightly larger, with a
few more bulbs.

_ Distribution.—Queensferry, Plymouth.

SCYPHOMEDUSAZA.
Order STAUROMEDUSZ&.
Family TESSERIDZ.

| Depastrum cyathiforme, (Sars 19).

_ Beaumont (5) found this species at Port Erin, in 1892,
and at Poyllvaaish, in 1893. It is fairly abundant in both
localities, attached to the under surface of stones near
the Laminarian zone.

Beaumont believes that Carduella cyathiformis, Allman,
Depastrum cyathiforme, Gosse, and Lucernaria cyath-
forms, Sars are one and the same species.

Heckel places Carduella cyathiformis in another genus,
as a synonym of Depastrella allmam, Heeckel.

Family LUcERNARID.

Haliclystus auricula (Rathke).
_A specimen taken by M. Chopin, in 1891, near Port
St. Mary. Mr. Beaumont has seen this specimen and

—_ iene ee NS SS es

984 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

believes it to be H. auricula, Rathke. Mr. Beaumont
(5) also gives a description of another species of Hali-
clystus, which may possibly be new.

Order DISCOMEDUS/AH.
Family PELAGIDA.
- Chrysaora isosceles, (Linnaeus 15).
Byerley (6) states that this medusa is rare, seen mostly
during July and August. Walker (7) states it is frequently
very common in the L.M.B.C. district.

Family CYANEID&.

Cyanea capillata, (Linnaeus 15). ;

Byerley (6) states that it is a very common medusa
during the summer.

I took a small specimen i in Port Erin Bay, on May
26th, 1893. Umbrella 14 inches in diameter. Larger
ones were seen at the beginning of June.

On April 30th, 1894, I captured a late Ephyra-stage —
showing the commencement of the long tentacles. On
May 9th, I caught a small adult specimen about 10 mm.
in diameter.

Family ULMARIDA..

Aurelia aurita, (Lamarck 15).

Aurelia aurita suddenly appeared in Port Erin Bay, on
June 2nd, 1893, and in a few days became abundant.
But throughout the whole of May I never saw a single
specimen. Diameter of the umbrella about 2 to 5 inches.

In 1894, on March 22nd, the first day of tow-netting, I
captured one specimen of the Ephyra-stage and a few at

_ the beginning of April. At the end of April a few of the

adult stage were taken; the largest 18 mm. in diameter.
The Hphyre are certainly scarce at Port Hrin and the

L.M.B.C. MEDUSA. WEste 985

large adults which are plentiful in the bay during the
summer probably come from far off breeding grounds.

Mr. Beaumont and I once dredged the scyphistoma-
stage near Bay Fine, August 1892; a few specimens on
an old Pecten shell.

Family PILEMIDA.

Pilema octopus, (Linnaeus 15).
Riuzostoma pulmo, Forbes (9).

During April, 1894, several large specimens of this
medusa were taken. Three were left by the tide on the
beach. Prof. Weiss found one which measured 23 inches
in diameter, and had arms 15 inchesin length. The margin
of the umbrella has a purplish border. On the external
side of the genital membrane many parasitic amphipods
(Hyperva galba) were found. This large medusa has been
recorded by Byerley. Mr. Walker states (7) that he has
seen many hundreds in a day.

REFERENCES.

1. Agassiz, A., 1865; ‘‘ North American Acalephe.”

2. Agassiz, L.., 1862; Monograph on Acalephe. Contrib.
Mareiaist., U.S: .Vol. LV.

8. Alder, J.,1861; Trans. Tynes. Field Club. Vol. V.,p.225.

4, Allman, G. J., 1871; Ray S. ‘“‘ Mon. Gymn. Hydroids,”’

5. Beaumont, W. I., 1893; Trans. Liverpool Biol. Soc.
Wareyil; p. 253; and “ Fauna,” Vol. IV.

6. Byerley, J., 18538; Liverpool Literary and Philosophi-
cal Soc. Vol. VIII., p. 125.

7. Clubb,J.A.,1886; Fauna of Liverp. Bay. Vol.I.,p.114.

8. Crawford, J. H., 1891; Ann. Mag. Nat. Hist., Ser. 6.
Wor. VILL., p. 296.

9. Forbes, H., 1848; ‘‘ British Naked-Eyed Meduse.”’

10. Garner, 1867; ‘‘ Holiday Excur. of a Naturalist,” p. 82.

286 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

11. Garstang, W., 1894; Journal Mar. Biol. Association,
Plymouth. Vol. III., p. 218.
12. Gosse, P. H., 1853; ‘‘ Devonshire Coast.” '

13. Green, J. R., 1857; Nat. Hist. Rev. Vol. IV., p. 242. —
14. Haddon, A. C., 1885; Fauna of Dublin Bay. Proc.
Roy. Irish Acad., Ser. 2. Vol. IV., p. 523.

15. Heckel, E., 1879; “‘ System der Medusen.”’
16. Hincks, T., 1868; ‘‘ British Hydroid Zoophytes.”
17. M‘Intosh, W. C., 1890; Notes from St. Andrews
| Laboratory. Ann. Mag. Nat. Hist. Ser. 6, Vol.
V., p- 296. =
18. Romanes,--G. J, 1876; Journ Linn. Soc. Zool.
Vol. XIT.; p. 524. Vol. XIII, p. 190, pl. XV—XVI_
19. Sars, M., 1846; Fauna Littoral. Norveg., Tom. I.
20. Sars, 1877; Fauna Littoral. Norveg., Tom. III.
21. Steenstrup, 1845; ‘‘ Alternations of Generations,”
Ray Soc.
22. Wright, T. S., 1858; Proc. Roy. Physical Soc., Edin.
no VOLS ap ar,

a,

re

287

[WORK FROM THE PORT ERIN BIOLOGICAL STATION, ]

REVISION of the AMPHIPODA of the L. M.B.C.
DISTRICT.
By AurreD QO. WatKeER, F.L.S.

With Plates XVIII. and XIX.
[Read April 5th, 1895.]

THE completion of that portion of Prof. G. O. Sars’
invaluable work on the Crustacea of Norway which
deals with the Amphipoda, has enabled me to name our
collection in accordance with it, and at the same time to
correct a good many errors that had occurred in our
former lists. As Prof. Sars’ work is likely to be the
standard for many years I have throughout adopted his
nomenclature for species described by him and would
refer any student of this class to it for descriptions. In
the case of species not described by him I have given
references to the best descriptions with which I am
acquainted. If I have occasionally ventured to differ as
to the validity of species with Prof. Sars I do so with the
full conviction that his knowledge of the Amphipoda is so
vastly greater than mine that in all probability he will prove
to be right. Many of these differences of opinion depend
upon the solution of the very difficult question whether
the Amphipoda, or some species of them, can become
sexually mature before they are full-grown. My own
opinion is that they can, and in support of this view I
have given various instances below of small females with —
ova. If this be proved (and it can only be done by rearing
them through successive stages in aquaria—a difficult and
generally impossible process) it will enormously increase

288 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the difficulty of defining species, but it ought to be
attempted at the Port Erin Biological Station. It might
probably be tried with success on Bathyporeia pilosa,
Lindstrom, (the form called B. pelagica, Bate, by Sars),
which is common in the harbour and appears to remain”
in shallow water. If young specimens were kept in:‘small
aquaria with sand at the bottom it would be seen what
changes they undergo. Very small females with ova were
dredged by Mr. I. C. Thompson, in June, in the harbour
and, if again taken, would form good subjects for the
experiment.

With regard to the new species described, the first,
Nannonyx spinimanus, is not only very distinct from the
only other species (N. goésiv) but might be thought to
require a new genus. It agrees, however, so well with
the rather peculiar external characteristics of Nannonyx— —
a genus so distinct that the points on which my species
differs might well be omitted and yet leave the definition
quite satisfactory—that I have preferred to include it, at
the same time pointing out the differences. On the other —
hand Amphilochus melanops and Photis pollex are so —
near to other species that were geographical sub-species —
_ permissible, I should have preferred to rank them as such.
I have given the lengths of the largest specimen of each

species in my possession for comparison with those of

other localities—the length including the uropods but not
the antenne. These measurements must, however, be
taken for what they are worth as it is impossible to be
certain that the specimens have reached their full size.

The specimens in the list have all been taken by dredg-

ing except where otherwise stated. The initials to some © a

of the species mean that the specimen was collected by
the gentlemen indicated, as follows : |

W.A.H., Prof. W. A. Herdman, F.R.S.

— ier
py

o

:

REVISION OF L.M.B.C. AMPHIPODA. 289

1.C.T., Mr. Isaac C. Thompson, F.L.S.

F.A., The late Frank Archer, Hsq.

The specimens from Galley Head, which is in the Irish
Channel but not in the L.M.B.C. district, were taken by
Mr. R. Li. Ascroft in trawl refuse or in a tow-net attached
to the trawl beam.

Of the species recorded the following are new to science,
viz., Nannonyx spinimanus, Amphilochus melanops, Pho-
tis pollex, and Podocerus herdmamn (P. odontonyx, Sars).
The following are, or were when I first published them,
new to the British Fauna: Orchomenella ciliata, Nan-
monyx goésr, Harpiua levis, H. crenulata, Metopa
bruzelu, M. pusilla, Paratylus falcatus, P. uncinatus,
Gammaropsis nana, Ischyrocerus minutus, and Siphon-
ecetes collettu; the last, however, has probably been
recorded as S. typicus, Kroyer which Sars considers to
be distinct.

It must be understood that the present list supersedes
all previous records of Amphipoda in the L.M.B.C. Reports
both as to species and localities.

In the notes on the species I make use of the following
abbreviations :— |

acc. app. = Accessory appendage of upper antenne;
ant.1=upper or first antenne; ant.2=lower or second

antenne; carp.=carpus, wrist, fourth joint ; dact. = dacty-

lus, finger, sixth and last joint; #.=flagellum of ant. ;
Gn.=first gnathopods, first pair of feet; Gn.?=second
enathopods, second pair of feet; Mdl.= Mandible; Maz.
=first maxille; Maz.?=second maxille; Maps.=maxil-
lipedes; ped. = peduncle; per. = pereon, mesosome ;
prp.’ = first to fifth pereeopods, third to seventh pairs of
feet ; pl.=pleon, metasome ; prop. =propodos, hand, fifth
joint; post.=posterior; seg.=segment; ¢.=telson; wp.
= first to third pair of uropods.

290 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

HYPERIIDA.

Hyperra galba (Montagu).
Females common in the bell of Rhizostoma pulmo
throughout the district: males rather scarce. Mr. R.
Newstead who collected some large ovigerous females
from lobster pots at Bull Bay, Anglesea, says that the
large eyes were very luminous. Length 10 mm.
Hyperoche tauriformis (Bate, 1868) =H. ree Boval-
lius, 1885.

One young male in a tow-net attached to a buoy at
Puffin Island. An examination of the remains of the
specimens labelled as above in Sp. Bate’s Collection, at
Plymouth, has satisfied me of the identity of the above
two species which were separated by Bovallius.* Bate’s
name of course should have priority. Length 3 mm.

[Parathemisto oblwia (Kroyer). |

A few taken in a tow-net attached to a trawl beam by
Mr. R. L. Ascroft, off Galley Head, Co. Cork. Length
5 mim. | |

ORCHESTIIDA.

Talitrus locusta (Pallas).

Abundant under drift-weed about high water mark, at
Colwyn Bay. Length1i8mm. ~—
Orchestia littorea (Mont.):=O. gammarellus ? (Pallas).

Under stones above high water mark. ‘Tal y Cafn,
Conway River; Burton Rock, Dee; Fleshwick Bay and
Port Erin, Isle of Man; at the latter place it is very
abundant and sometimes invades the laboratory in great
numbers. One specimen taken at dead low water at
Colwyn Bay. Length 18 mm. :

Hyale milssonw (Rathke).

Among stones and alge on shore at Hilbre Island, ae

* Monograph of the Amphipoda Hyperiidea, pt. IL., p. 445,

-- -- REVISION OF L.M.B.C. AMPHIPODA. 291°

Puffin Island, Porthwen Bay, Anglesea, and Port St.
Mary. One young 10 to 17m. N.W. of Mersey Bar 10 to
14 fath. 27/9/90 (1.C.T.). Length 5 mm.

LYSIANASSIDA.
Lysianax longicornis (Lucas) =L. ceratinus,* Walker. »
Little Orme; Puffin Island; Menai Straits; 6 to 12
fath.- I have never taken a male with fully developed
lower antenne. An old female taken off the Little Orme
had these organs rudimentary and the eyes imperfectly
developed. Colour dull yellow, eyes large, dark. Length
9mm.
-Socarnes erythrophthalmus, Robertson. +
Port Erin 15 to 20 fath., 24/3/94; Menai Bridges 5 to
7f. Common April and May, 1894. Length 44 mm.
Perrverella audowwmiana (Bate) = Lysianaz audowwmana,
Bate.
22m. 8.H. Isle ot Man, 21/5/88, 30 fath. on sponge;
on Pecten maximus, 1/4/93 (I.C.T.); Menai Bridges,
2/4/94, 5 to 12 f. rocky ground; off Fleshwick Bay, 8/7/94,
broken shells and gravel, Pectens, &c., 22 f. Having
succeeded in cleaning the type specimen, in Bate’s
Collection at the British Museum, I have no hesitation
in saying that it is this species.t Length 24 mm.
Callisoma crenata, Bate.
7m. W. of Bradda Head, 31 f., 25/4/95. One female.
[One male with adult antenne, length 4 mm., from trawl
off Galley Head, Co. Cork, 24/10/94 (R.L.A.)}. Length
4 mm.
Hippomedon denticulatus (Bate).

* Fauna of Liverpool Bay, Vol. Il. Third Report on Higher Crustacea.

+Second List of Amphipoda and Isopoda of the Firth of Clyde, &c.
Trans. Nat. Hist. Soc, Glasgow, Vol. III., p. 200, 1892; also Bonnier,
Bull. Sci. de France, Vol. XXIV., p. 183, Pl. VI.

¢ Bonnier, l.c., p. 181, Pl. V.

292. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

- Port Erin, electric light, 20/5/88. One young. [Galley
Head, one adult male. Length 14 mm.] be

Orchomenella ciliata, Sars=Tryphosa ciliata, Sars.

Between Isle of Man and Liverpool, 30 fath.; Puffin
Island, 15 f.; Colwyn Bay, 24 f., 1/2/90; off Port Erin,
15 to 20 f., 24/3/94. [Galley Head, one male. Length
33 mm. | '

Nannonyx goéswt (Boeck) = Orchomene goésu, Boeck.

Puffin Island, low water spring tide, 8/8/88. Be
2+ mm.

N. spinimanus, n.sp.* (Pl. XVIIL., figs. 1—11).

Menai Bridges, 2/4/94 and 31/5/94, rocky bottom,
5 to 8 f. |

This species falls into the genus Nannonyz, G. O. Sars,
except as to the maxille, maxillipedes, gnathopoda and
pereiopoda. In these there are more or less important
differences but not sufficient in my opinion to constitute
a new genus. |

Body short and thick. Head as long as first segment — 4

of per. lateral angles produced and rounded. Anterior
coxal plates about as deep as the body, angles rounded.
Third pl. seg. with hind margin straight, serrate, hinder
angle rather acute with the point blunt; fourth seg. ys
a prominent rounded hump, fifth very det! }

Ant.! with first joint of ped. longer than the two next, fl.
seven-jointed, the first as long as the remaining six,
second short and wide, acc. app. four-jointed the first
longer than the remaining three and as long as first jou 7
of fl. which is very hairy.

Ant.? slender, fl. in both sexes five-jointed and shorter
than ped. |

* Walker, A. O., Rep. on Marine Zool. of Irish Sea. Brit. Ass., 1894, : a ‘.

p- 327.

REVISION OF L.M.B.C. AMPHIPODA. 293

Mandibles long ; cutting edge longer than in N. goési,
yellow and strongly refractive.

Max.! with outer lobe much longer and broader than
inner which has two sete at the tip. ©

Max.” with the inner lobe twice as wide as the outer.

Mxp. with the base moderate sized, not setose; the
masticatory lobe large and armed with strong triangular
tooth-like spines at the tip and nearly half way down the
inner side, reaching to the middle of the second joint of
_ the palp; basal lobe narrow and only half as long as the
masticatory; palp with third joint longer than first or
second, last joint of the usual claw form. Eyes very large,
oval, dark brown. .

Gn.' Strong, first jomt nearly as long as last three;
carp. more than half the length of prop.; prop. in male
tapering to base of dact., armed with five or six short
stout spines along the hinder margin; the female has the
margins almost parallel, the postero-distal extremity
rounded and provided with three spines placed close
together, not setose; dact. rather short with three spinules
close together in the middle of the hind margin.

Gn.* Prop. subchelate, rather shorter and nearly as
wide as carp. :

Prp. First and second with first joint about the same
width as third (merus) which is longer than fourth, and
about the same length as fifth; the last three prp. increase
in length successively but slightly; first joints dilated,
hind margins entire; the third joint in third and fourth
prp. 1s wider and more produced posteriorly than in the
fifth ; dact. of moderate size.

Up. First and second with rami shorter than ped.
spinous; third with inner ramus much shorter than
outer, styliform ; in the male the outer is rather shorter
than the ped. and furnished at the tip with a tuft of long

SSS

294 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

plumose sete; in the female the outer is about half as
long as the ped. very stout and with two short spines
near the base of the nail, the inner ovate with two or
three minute spinules at the tip and one spine lower down.

Telson erect, convex, as broad as long, distally rounded
with two spines on the outer margin, and a setule behind
each. Colour brown. Length 44 mm. Easily distin-
guished from N. goésw by the longer, more rectangular,
glabrous and spinous prop. of first gnathopoda and by the
smaller coxal plates of the per. Four specimens were
taken in the deep holes near the Menai Bridges on a very
rocky bottom. 7 haa

Tryphosa nana (Kréyer) = T. horringu, in third Report

on Higher Crust. |

Puffin Island, 10 to 15 f., 24/3/88; Menai Bridges,
31/5/94. Length 4 mm.

Tryphosa horringu, Boeck.

Bull Bay, Anglesea, ‘‘from ambulacral grooves of
common Starfish” (F. A.).

[Tryphosites longipes (Bate).

Galley Head, two males. Length 14 ma.

Hoplonyx similis, Sars.

Laxey Bay, 8 fathoms, 24/9/92, one sp. The 0 eyes
lost their colour in‘'a mixture of meth. spirit, glycerine,

and water in a few days, becoming very pale yellow.
Length 10 mm.

[Lepidepecrewm carinatum, Bate.

Galley Head, two males, 7 mm. |

Euonyx chelatus, Norman. |

Puffin Island, off the lighthouse; between Holyhead
and Isle of Man, 50 f. on Hchinus sphera 20/7/89; 10 to
17 m. N.W. of Mersey Bar on Hchinus 27/9/90; hens 2
8 mm.

REVISION OF L.M.B.C. AMPHIPODA.. 295

PONTOPOREIID.
Bathyporeia norvegica, Sars.
Lianfairfechan, dug out of sand between tide marks
1/9/83; Port Erin, electric ight; Colwyn Bay; off Gar-

wick Head, Isle of Man 24/9/92. Length 8} mm. A

small one from the same locality measuring 44 mm. had
the characteristic acute and up-turned post. angle of the
third pleon seg.

Bathyporeia pelagica, Bate = B. pilosa, Lind.,in Rep. ITI.

Port Erin Harbour, abundant; Colwyn Bay; Llanfair-
fechan; Menai Straits. Length of ovigerous females from
2mm, to 44 mm. Males with fully developed antenne
from 3 mm. to 44 mm. from the same locality.

This is the commonest form of what with all deference
to Prof. Sars’ superior knowledge I cannot but consider
with Mr. Stebbing to be but one species, viz., B. pilosa,
Lindstrom. The differences that Sars indicates between
B. pelagica, B. robertsonu, B. gracilis and B. pilosa
depending mainly on the colour of the eyes and the
presence or absence of the small spines on the first urus-
segment appear to me insufficient for specific distinction.
I have known the red eyes of B. pelagica turn white in
some specimens (and not in others) after two days in weak
spirit and glycerine while in other specimens they become
dark. The eyes of the specimen marked B. pelagica in the
Sp. Bate Collection in the British Museum are large and
dark—this is an adult male. In the same collection are
eleven specimens in one tube marked B. pilosa (Lind.)
some of which have, and others have not the spines on
the first urus-segment. One large female has a rudimen-
tary tooth on the lower margin of the third pleon segment.
B. pelagica (or pilosa) becomes sexually mature at a very
early period, specimens only two mm. long containing one

to three ova having been dredged by I. C. Thompson on

June 19, 1892.

296 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Eionaurtns arenarius (Slabber).

Llanfairfechan, shore; Colwyn Bay, shore 31/7/90,
Length 8 mm.

Urothoé brevicornis, Bate. . ,

Lilanfairfechan, shore, Sept., 1883. Dug out of sand
between tide marks. Length 6 mm.

Urothoé elegans, Bate = U. marinus, Rep. IIL. , p. 205.

Puffin Island . C. T.); Menai Straits bere Beau-
maris and Garth 5 to 10 f., 16/5/91 and 17/9/94; Little
Orme 4 to 7 fathoms, 14/9/94; length 3} mm. The above
two species are very difficult to separate and it is possible
that the latter is only the young of the former. - Mr.
Stebbing has treated the genus exhaustively in Trans.
Zool. Soc. (uondon), Vol. XTIL., part 1, 1891, Pls. I—IV.

Urothoé marinus, Bate.

2, miles §.E. of Kitterland 17 f., 27/5/94.

Several specimens including adult male and female:
these have been examined by Mr. Stebbing who considers
them to be this species. Length 5 mm.

. PHOXOCEPHALIDZ.

, Phoaocephalus fultom, T. Scott, (see Highth Ann. Rep.
of Fishery Board, Scot., 1890, p. 327, Pls. XII., and
XIII., for female and young-male; and Robertson,
D., second list of Amphipoda, &c., of Firth of Clyde,

for adult male) =P. chelatus Delle Valle, Gammarini
of the Gulf of Naples.

Port Erin, electric light, 1888 and dredged 15 to: 200%
24/3/94; Menai Bridges 5 to 12 f., 30/5/94; length 23 mm_
I have also taken it off Jersey and Guernsey. The large
eyes would seem to place this species in Paraphoxus, Sars,
but the gnathopoda are distinctly unequal.

Paraphozus oculatus, Sars.

7 miles W. of Bradda Head 31 £. Ataion one ae

aS)

REVISION OF L.M.B.C. AMPHIPODA. 297

Harpinia neglecta, Sars=H. plumosa (Kr.), in Rep. III.

Between Isle of Man and Mersey Bar, 10 to 20 f.,
3/9/87; Colwyn Bay, 24 f.; Menai Straits, 16/5/91; 5 m.
W. of Dalby, Isle of Man, 30 f. sandy mud, 8/7/94.
Length 3} mm.

None of the specimens had the pleon hairy as described
by Sars, and as the character of the post antennal angle
is very difficult to see I have great hesitation in deter-
mining whether they should be referred to this species or
to H. plumosa (Kroyer). As, however, the latter appears
not to have been met with south of the Arctic circle it is
safer to call them H. neglecta.

Harpina crenulata, Boeck.

7 m. N.W. of Bradda Head, Port Erin, 39 f., 29/1/93.
Three adults and one young. Length 24 mm.

Harpima levis, Sars.

7m. W. of Niarbyl, Isle of Man, 45 f., mud, 8/7/94.
Length 25 mm.

AMPELISCIDA.

Ampelisca typica, Bate=A. tenwicornis, Rep. III., 207.
Port Erin, electric light, 21/4/89. Length 93 mm.
Ampelisca tenuicornis, Lilljeborg.

Port Erin; 5m. W. of Dalby, 30 f., sandy mud, and 6
m. W. of Contrary Head, 38 f., mud, Isle of Man, 8/7/94.
Length 9 mm. .

Ampelisca brevicorms (Costa)=A. levigata, Lilljeborg.

Between Isle of Man and Orme’s Head, 20 to 380 f.,
28/8/86; Colwyn Bay, 3 f., 24/5/87; Bull Bay; Port
Erin—electric light; off Southport, 10 to 20 f., June ’91.
Kyes crimson with a scarlet line behind them and five
black stellate spots behind that. Lower part of head
having a scarlet cloud extending to the first epimere.
Remainder of body transparent white with scattered black
stellate spots. Length 13 mm.

ro
1 ona
A

298 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. —

Ampelisca spinipes, Boeck = A. tenuicornis, Rep. III., 207.
Throughout the L.M.B.C. district in 20 to 50 fath.
Length17mm. This is the commonest species in the dis-
trict, the preceding one being the next commonest. Ihave

little doubt that the species figured as A. gawumardw (Kr.)in

the British Sess.-eyed Crust. is this species and not as Sars
supposes, A. typica (Bate). Ihave examined Bate’s speci-
men, in the British Museum, and find both it and the figure
to confirm this view. The relative proportions of the upper
and lower antenne which are correctly drawn, are alone
sufficient to show that it cannot be A. typica.

Ampelisca macrocephala, Lilljeborg. —

Port Hrin, Aug., 1893 (W. A. H.).. Juength = 20}
Resembles A. brevicornis in the form of the hind margin
of the third pleon segment, but may be distinguished most
readily by the greater length of the upper antenne, and
by a long spine near the tip of the outer ramus of the
second uropods. )

Haploops tubicola, Lillje.

One specimen between Isle of Man and Orme’s Head,
20 to 80 f., 28/8/86. Length 8 mm.

AMPHILOCHIDA.

Amphilochus manudens, Bate. _

Little Orme, Aug. and Sept., 89: Gt. Orme, 8 f.,
1/4/90; Puffin Island; Menai Bridge, 5 to 12 f., April and
May, 94; 2 m. S.H. of Kitterland, Isle of Man, 17 £3
27/5/94. Colour generally brownish, sometimes almost
black; one specimen from Gt. Orme bright scarlet. Eyes
red, rather small. Length 23 mm.

Amphilochus melanops, n.sp.* (Pl. XVIIL., fig. 12; PL.
NS ies 13 No):

Little Orme, 5 to 7 f.; Menai Straits near Beaumaris,
5 to, 10 £5. 17/9/94.

* Walker, Rep. on Mar. Zool. of Irish Sea, Brit. Assn., 1898, p. 535.

REVISION OF L.M.B.C. AMPHIPODA. 299

Head much curved, about as long as the two first per.
segs., lateral angle rounded. Eye round with large dark
brown centre. First coxal plate small oval, second
rounded at lower margin, which is minutely crenate;
third less deeply crenate; fourth and fifth entire or
smooth; per. segs. increasing successively in length, the
last equal to the first pl. seg. Pl. segs. all somewhat
produced but rounded at the post. angle. Ant.! in female
shorter than ant.’, fl. 7—8 jointed as long as ped.; ant.’
with last joint of ped. longer than preceding, a small tooth
on the lower margin of the distal end of second and third
joints; fi. slender 7—8 jointed shorter than ped.

Gn.’ much smaller than en.?, first jomt as long as the
next four combined, merus and carp. having the ends
truncate and setose the carp. prolonged to half the length
of the post. margin of the prop., anterior margin of prop.
slightly convex and not produced into a tooth. Dact.
serrate on the proximal two-thirds of its length the
serration ending in a secondary tooth.

Gn.” of the same general form as gn.’ but the carpal
process reaches almost to the post. angle of the palm;
the anterior margin of the prop. is slightly concave.

Prps. There is nothing distinctive about these.

Up. First “‘ eque-attinent,” (¢.e., reaching as far back)
as the third, extremity of longest ramus of up.” not quite
reaching to the end. of ped. of up.*. It is rare to find a
specimen which has not lost its third uropods.

Telson concave forming a triangle of which the sides
are little longer than the base reaching about one-third of
the length of the ped. of up.®
_ This species is undoubtedly very near both to A.

marionis, Stebbing (‘‘ Challenger’? Amphipoda, 1888), and
to A. brunneus, Della Valle (Gammarini, Fauna des
Golfes v. Neapel, 1894). From the former it differs in its

SS

Seo a _
SANT AE AIF TT EAE ICI ta LON ABE TE Bee Aa Si ae Sit re.

; f
i
ih
i
i
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i

——— — -~ —_
3 a
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RE = SN A Wg ER

300 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

conspicuously large and dark eyes and in the less convex
palm and greater relative length of the prop. of the
gnathopods. From the latter the principal difference is
in the much shorter telson. The eyes also differentiate
it at once from A. tenwimanus, Boeck, and from A.
manudens, (Bate), with which I have found it associated. -
From this species it is also distinguished by the rounded
lateral angle of the head, and by (as also from A. ocwlatus,
Hansen) the absence of the distal tooth on the anterior
margin of the prop. of the gnathopods. Males of this
species are scarce. Colour brown. Length 23 mm.

Amphilechoides odontonyx (Boeck). )

8m. W. of Fleshwick Bay, 33 f., 5/6/92. Length 2mm.

Gitana sarsiw, Boeck.

Little Orme; Menai Straits, 5 to 10 f., 17/9/94; 8 m.
W. of Fleshwick Bay, 33 f. Length 24 mm.

Cyproidia brevirostris, T. and A. Scott (Ann. and Mag.

N. H., 18938, Vol. XII.,p. 244, Pl. X11)

8m. W. of Fleshwick Bay, 33 f. Length2mm. Very
near C. damnoniensis (Stebbing) from which its most
obvious difference is the concave lower margin of the first —
joint of the last pereopods. In C. damnoniensis the
lower margin is convex. |

a

STENOTHOIDA.

Stenothoe marina (Bate).

Common from the mouth of the Dee to Menai Straits,
2to 15 f.; off Fleetwood and Blackcombe. ‘Two speci-
mens 6 m. §.E. Calf of Man, 34 f., 25/4/95. Colour white
or stained with red especially about the head. Length4mm.

Stenothoe monoculoides (Montagu). 3

Abundant in tidal pools Port Erin, Fleshwick Bay, and
Port St. Mary, Isle of Man; Menai Straits, April and May,
94, 7 to 12 fath., rocky ground, common. Length 33} mm.

REVISION OF L.M.B.C. AMPHIPODA. 301

This species seems to prefer rocky ground while S. marina
is found on sand, I have not met with it HE. of Menai
Straits.

Metopa alderi (Bate).

Menai Bridge, April and May, 94, 7 to 12 f.; Turbot
Hole, Puffin Island. Colour white with red blotches on
second, third, and fourth epimeres and first jot of cor-
responding legs, also on the urus. Length 45 mm.

Metopa borealis, Sars.

Little Orme, 5 to 10 f., common; Menai Straits; Puffin
Island, 14 f. Colour uniform white or grey. Length
2mm.

Metopa pusilla, Sars.

Rhos Bay just below tide mark, 18/5/93; Menai Bridge,
7 to 12 f., 2/4/94. White, tinged with red on the back
and limbs. Length 2 mm.

Metopa rubro-vittata, Sars.

Little Orme, 4 to 7 f.; Colwyn Bay, 24 f.; Menai
Bridge. One specimen was beautifully and uniformly
speckled with bright red. This species 1s best distinguished
by the spine on the posterior margin of the propodos of
the first gnathopods. Length 24 mm.

Metopa bruzeli (Goés).

Colwyn Bay and Little Orme, common; off Port Erin,
geet 0/0/92; Port Erin harbour, Nov. ’92 d.C.T.).
Colour white with large blotches of brillant red on front
segments and epimeres. Length 27 mm.

Cressa dubia (Bate).

Great and Little Ormes; Colwyn Bay; Menai Straits ;
10 to 17 m. N.W. of Mersey Bar; 2 m. 8.E. of Kitter-
land, Isle of Man, 27/5/94, 17 fath. Colour pale yellow
clouded with red on fifth, sixth, and seventh segments of
pereon. Length 2 mm,

302 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY. —

LEUCOTHOIDA.

Leucothoe spinicarpa (Abildgaard).

S.E. coast of Isle of Man, in branchial sac of Ascidia
venosa and A. mentula; off Port Erin, washed out of
Pectens (I. C. T.); Towyn, Anglesea, 5f. Length 12 mm.
Generally speaking only one or two individuals are found
in each Ascidian.

Leucothoe lulyeborgu, Boeck =? L. amparicorms, Nor.

Off Port Erin, ‘‘ Lady Loch,” 24/3/94, ‘“‘ washed out of
dredged material from several hauls” (1 .C. T.).’. Length
94mm. Of the above two species the latter (indicated by
the up-turned hinder angle of the third pleon segment and
other characters) had the upper and lower antenne of the
same length, while in the former the upper were much
longer than the lower, showing how fallacious characters
derived from the relative length of these organs are. My
specimen was doubtless immature.

AMDICERIDA.

Monoculodes carinatus, Bate.

Port Erin, outside harbour, 21/8/92 (I. C. T.) [Galley
Head, 24/10/94] 7. m. W. of Bradda Head, 31 f., 25/4/95.
Length 3 mm. ' :

Perwoculodes longimanus (Bate).

Port Erin Harbour; Colwyn Bay; Menai Straits. —

Shore to 89 fathoms, sandy ground. Eyes generally red
but sometimes dark. An abundant species where it occurs.
Length 4 mm.

Pontocrates arenarius (Bate) =P. norvegicus in Rep. II.

Rhos and Colwyn Bays; Menai Straits; Port Erin
- Harbour, common; Garwick Head; in stomach of Agonus,
Morecambe Bay; a female 6 mm. long, 1 m. 8.EH. of
Kitterland 20 f., average length of females with ova from
Port Evin Harbour 3 to 4mm. Colour white,

REVISION OF L.M.B.C. AMPHIPODA. 303

Synchelidium haplocheles (Grube, not G. O. Sars)=S.

brevicarpum, Sars.

Tittle Orme; Port Erin and Ramsey Harbours at
electric light; off Port Hyrin, “John Fell,” 8/7/94. 22 f.
Length 3mm. The dark markings on the body segments
is the most easily seen distinction between this species
and the preceding.

PARAMPHITHOIDA.

Paramphithoé bicuspis (Kroyer).

Dee to Menai Straits, shore to 17 f.; between Holyhead
and Isle of Man 40 to 60 f., 20/7/89; Towyn, Anglesea ;
10 to 17 m. N.W. of Mersey Bar, 27/9/90. Length 12 mm.

A common species on the N. coast of Wales but not
met with hitherto on the Isle of Man coast. It is variable
in colour, generally closely freckled with brown but some-
times almost pure white. I cannot regard P. monocuspis,
Sars, as anything but the young of this species, as I
_ have constantly taken them together and always observed
that only the largest specimens have a dorsal tooth on the
first pleon segment and this varies in length in proportion
to the size of the specimen.

Paramphithoe assvmilis, Sars. (Pleustes glaber, Rep.

IV., p. 241.)

Puffin Island, shore; Great and Little Orme’s Heads;
10 to 17 m. N.W. off Mersey Bar; Calf Sound; Menai
Straits 10 to 12 f., 2/4/94; off Blackcombe. Colour ereen-
ish white sparingly mottled with brown. Length 5} mm.

Stenopleustes nodifer, Sars.

Little Orme and Rhos Bay, 4 to 7 f., April and July,
1893; 7 miles W. of Bradda Head, 31 f., 25/4/95.

The “nodiform projections” in the specimens taken
were reduced to a mere emargination of the hind margin
of the two anterior segments of the pleon. The limbs

304 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and greater part of the body are speckled with dark red
of which colour there is a dark cloud on the fourth, fifth
and sixth pereon segments. The immense reniform eyes
are very characteristic. Length3 mm. _

KPIMERIDA.

[pumeria cornigera (Fabricius).
Galley Head, 24/10/94. uength 10 mm.]

- J[PHIMEDIIDA.

Iphimedia obesa, Rathke.

Throughout the district 2 to 15 f. Length of large
female taken about 17 miles N.W. of Mersey Bar,
24/9/90) 5 salir

Iphimedia nunuta, Sars.

Colwyn Bay, 22/1/92, &c. ; Little Orme, 14/9/94; Menai
Straits, 17/9/94, 10 to 12 f., abundant; Port Erin, 15 to
20 f., 24/3/94. Generally lighter coloured than I. obesa,

sometimes almost colourless (Menai Straits). Length

Syiaau gan

LAPHYSTIIDA.

Laphystius sturioms, Kroyer.

One specimen from underneath the pectoral fin of a
Cod from Liverpool Bay (Lancashire Fisheries Labora- _
tories, November, 18938). Liength 8 mm.

SYRRHOIDA.

Syrrhoe fimbriatus, Stebbing and Robertson (Trans.
Zool. Soc., London. Vol. XIIL., part 1, 1891, p. 3l,
ey):

Two miles §.E. of Kitterland, Isle of Man, 17 fathoms

27/5/94 (I. C. T.); 7 miles W. of Bradda Head, 31 f.,
25/4/95; four specimens. Length of female with ova
12 mm.

ta.
1 yeas

REVISION OF L.M.B.C. AMPHIPODA. 305

EUSIRIDZ.

Husirus longipes, Boeck.
Off Port Erin, August, 1893 (W. A. H.). Length 5} mm.

CALLIOPIIDA.

Apherusa bispinosa (Bate).

In tidal pools and shallow water throughout the district
generally abundant. Very variable in colour—sometimes
black. Length 6 mm. A small female with ova meas-
ured only 33 mm.

Apherusa jurinn (M. Edwards) = Calliopius norvegicus,

(Rathke).

Common in tidal pools throughout the district. Colour
whitish mottled with red.. Length 8 mm.

Calliopius leviusculus (Kroyer).

Another common shore species. Generally greenish
white and without markings but sometimes mottled with
red. It is a question whether C. rathket (Zaddack) is
not the young of this species. A large female with ova
from Colwyn Bay measures 13 mm., another only 64 mm.;
no doubt Sars would call this C. rathket, which form was
common at Puffin Island in tidal pools September 9, 1888.
Tt differed from the commonest form of C. leviwsculus in
being spotted with red.

ATYLIDZ.

Paratylus swammerdamu (M. Edwards).

Abundant throughout the district, shore to 20 fathoms.
Adult males and females reach 9} mm. in length but
males with the ciliated antenne characteristic of sexual
perfection and females with ova, both only 4 mm. long,
were taken in tidal pools at Colwyn Bay and in 4 to 7 f.
off the Little Orme in July, 18938. One of the small
females had young ones escaping from the incubatory
pouches,

806 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Paratylus faleatus (Metzger). | | 4

Little Orme; Colwyn Bay. White with ared spoton
the back of each segment except the fourth pleon. Length .
5 mm. |

Paratylus uncinatus (Sars, Oversigtaf Norg.Crust.,p.102),

Red Wharf Bay, Anglesea, 20 f., 8/6/89. This species
which has been accidentally united by Sars in his Norwe-
gian Amphipoda with the preceding, differs from it only
in not having the pleon segments furnished with dorsal
teeth. The species figured in the above work is this form
and not Metzger’s. The difference does not appear to be
one of age or sex. Length about 5 mm.

Paratylus vedlomensis (Bate).

Puffin Island—low water; Port Erin Harbour, electric a

light, &c.; 8 miles W. of Fleshwick Bay, 33 fathoms; 3
miles W. of Calf of Man, 19 f. (Galley Head). Length
8 mm.
Deaxamine spinosa (Montagu). 3
Common throughout the district; shore to 10 fathoms. — 4
Length 15 mm. ;
Dexamine thea, Boeck. |
Port Erin Harbour, 4/6/92, several. Length 44 mm.
Triteta gibbosa (Bate). | _
Colwyn Bay; Anglesea Coast; Puffin Island and Port
Erin encysted in the outer integument of Ascidians; 16
miles N. of Holyhead 40 to 50 fathoms; Menai Bridge
very abundant; adult males at electric ight, Port Erin,
21/4/89. Length5mm. It is remarkable that the emar-
gination of the ant. margin of prop. of the first gnath. in
the adult males of this species which caused Nebeski to

make a distinct species of it (I. dolichonyz) should have 4

escaped the notice of so many carcinologists, including
even so careful and accurate an observer as Prof.G.O. |
Sars. Canon A. M, Norman informs me that such males

=

—

REVISION OF L.M.B.C. AMPHIPODA. 307

occur ‘“‘among his Shetland specimens examined by
Spence Bate when engaged on his work.”’
Guernea coalita (Norman, Ann. and Mag. N. H., Ser. 4,
Vol. IT., 1868). ,
Off Great Orme’s Head, 1/4/90; 25 miles 8.E. of Calf,
6/6/92; Off Port Erin 15 to 20 f., 24/3/94; 2 miles 8.H.
of Kitterland, 17 f., 27/5/94. Length 13 mm.

GAMMARIDZ.*

Melphidippella macera (Norman).

8 miles W. of Fleshwick Bay, 33 f., 5/6/92; 7 miles W.
of Bradda Head, 31 f., 25/4/95. Length 5 mm.

Amathilla homari (Fabricius) =A mathilla sabini (Leach).

The young of this species is one of the commonest
Amphipoda on our coasts, in tidal pools during spring and
earlysummer. I cannot doubt that A. angulosa (Rathke)
is this young form. The large females measuring 26 mm.
in length come to the shore in early spring to deposit their
young and may then be taken among Algz. The young
at first have no dorsal carina which is only developed by
degrees; in their earliest stage these are Grayia wmbricata
(Bate). The largest I have of the young form is 9 mm.
long, nor have I have ever seen one between these sizes.

Gammarus marinus, Leach.

Puffin Island; Hilbre Swash; Colwyn Bay; Port Erin
Harbour. Length 16 mm.

Gammarus locusta (Linné).

Common everywhere on the shore under stones between
tide marks. Length 25 mm.

Gammarus pulex (De Geer).

In brooks and springs up to 700 feet above the sea.
Length 16 mm.

* For the Gammaride see a valuable paper by Canon A. M. Norman in
Ann. and Mag. N. H., Ser. 6, Vol. 4 (1889).

“ Se pa emt ee
“ a "al ae ba

808 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Melita palmata (Montagu).

Colwyn Bay; Puffin Island; Port Erin; shore to 30 t
Length, excl. of third uropods, 9 mm.

Melita obtusata (Mont.).

Throughout the district: sometimes on Echinoderms,
shore to 30 f. Length, excl. third uropods, 9 mm. 7

Mera othonis (M. Edw.)=M. longimana [second Rep.

L.M.B.C.] (Bate and West., male) M. othoms (Bate
and West., female).

Bull Bay, Anglesea, 20 f., 8/6/89; off Port Erin and
Fleshwick Bay, 15 to 24 f. Colourless or light brown.
Length, excl. of third uropods, 12 mm.

Mera semi-serrata, Bate.

Off Port Erin, 15 to 25 f., 5/6/92; 24/3/94. Light brown.
Length 5 mm.

The fact of the above two species being found together _
| and their resemblance in many points, coupled with the
smaller size of M. semi-serrata, would seen to point to
this species being the young of M. othonis. In my
specimens of the latter the lower margin only of the third —
pleon segment is serrated, while in M. semi-serrata it is —
| the hind margin which is so. But Bate and Westwood
| | in their figure and description of M. othonis (female) show 7
| both margins serrate. This seems to indicate an inter- a
mediate age when the one form is passing into the other.
Mera bater, Norman (Ann. and Mag. N. H., Ser. 4,
Voli yePr Os):
M. Hainan (Bate and West., Vol. Il., p. oy
male).
Gammarus tenwwmanus (Bate and West., Vol. 1,
p. 384, female; Bate, Cat. Amph. Brit. Mus., 7
p. 214). a
Puffin Ieland, 15 f., April, 81; on Spatangus, 3 m. off .
Dulas Bay, 14 f., 8/6/89; Menai Straits, 10 to 15 £9 4

REVISION OF L.M.B.C. AMPHIPODA. 309

17/9/94; off Port Erin, 15 to 20 f., 24/3/94; 10 to 17 m.
Ney. of Mersey Bar. Length6mm.

I have examined the specimen marked Gammarus
tenuimanus, in the Sp. Bate Collection at the British
Museum, and am satisfied it is this species. The des-
cription and drawing of the second gnathopod in the Brit.
Sess. Hyed Crust. are very faulty; the description in the
catalogue is much more accurate.

Megaluropus agilis, Norman (Ann. and Mag. N. H.,

Ser. 6 (1889), Vol. III., p. 446, Pl. XVIIT.).

Common on the N. coast of Wales and the Isle of Man,
2 to 10 fath. Colouring when alive very beautiful. It
has a bright crimson blotch on the head above the eye;
telson and base of third uropods yellow—the latter with
three or four opaque white spots on the outer edge of the
outer ramus. Length 4 mm.

Chetrocratus sundevalli (Rathke).

Not uncommon on the N. coast of Wales and the Isle
of Man, from 2 to 30 fath. Margins of the first joint of
all the legs and both ant. scarlet; segments and epimeres
spotted with the same colour. Length 8 mm.

Chevrocratus assvmilis (Lilljeborg).

Port Erin harbour, 28/1/93; 7m. W. of Bradda Head,
81 f., 25/4/95. Colourless, by which, when fresh, the
female may most readily be distinguished from the pre-
ceding species. See also Norman in Ann. and Mag.
Weer oer, 6 (1889) Vol. [V., p..1381, Pl. XI. and XII.
Length 7 mm.

Lilljeborgia pallida Bate.

Bull Bay, Anglesea, 8/6/89, 17 f.; Port Erin, 22/8/98
(W.A.H.). Length 44 mm.

Lilljeborgia kinahani (Bate).

38m. W. of Calf of Man, 29/1/93; Menai Straits off
Rhianfa, 30/5/94, 7 f. Length 3; mm.

:= = —_
ot eae
a i

310 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

PHOTIDZ.

Aora gracilis (Bate).

North coast of Wales, 2 to 15 fath., rather common ;
Port Erin; (Lambay Isle,28/10/94, R.L.A.]. Length8mm.

Autonoe longipes (Lilljeborg). .

Anglesea coast, “‘ Spindrift,” 8/6/89, one male ; Laxey
Bay, 4 to 12 fath., 24/9/92; 5 m. W. of Dalby, 30 f.,
8/7/94; Calf Sound, 30/8/94: Menai Straits. Length 3 mm.

Leptochetrus pilosus, Zaddach.

Protomedeia pectinata, Norman (Final. ner of Bi

Shetland Dredging, 1868).

Protomedeia pilosa (Zadd.) and P. hirsutumana —

Bate? Grube, Beitr. z. Kenntniss der Istris-
chen Amphipodenfauna.
Leptocheirus pilosus (Zadd.) Della Valle, Fauna
des Golfes v. Neapel—Gammarini. , :
Menai Bridge, April and May, 94; washed out of 4
dredged stuff, off Port Hrin, many young, 24/3/94 (1.C.T) ; 4
2m. 8. EH. of Kitterland, 17 f., 27/5/94. Colour deep
yellow with transverse brown lines on all the segments.
Length of female with ova 2} mm.
_Leptocheirus hirsutumanus (Bate).
Protomedia hirsutimana, Bate.
Leptochewus ptlosus (Sars, not Zaddach).
One specimen 2 m.‘8. E. of Kitterland, 17 f., 27/5/94.

Colour very pale yellow without markings. Length 53 mm. — 4

I regret that I am unable to agree with Prof. Sars in q
assigning the species described by him, and which he _

rightly identifies with P. hirsutimana, Bate, to L. ptilosus, =
Zaddach. The latter author in describing his species

entirely overlooked the secondary appendage, which, —
however, Fr. Muller subsequently stated to be one-jointed. 4

This Zaddach could hardly have done had it been the | 4

long six-jointed appendage of Sars’ species. In JL.

REVISION OF L.M.B.C. AMPHIPODA. Bile

pectinatus (Norman) which I consider identical with L.
pilosus (Zadd.) this appendage is two-jointed and very
small and the minute second joint might easily be over-
looked by Muller, as indeed might the whole appendage
by Zaddach. Again, the form of the propodos of the first
gnathopods, described by Zaddach as somewhat swollen
and broader towards the extremity, does not agree with
Sars’ figure; and the dactylus of the second gnathopods
which is described by Zaddach as straight and feeble
(“...ungue recto debilique terminatur’’) and which in
L. pectinatus is a very characteristic thin narrow lamina
terminated by a small tuft of sete, is quiteenormal in L.
hirsutumanus. Finally, Zaddach could hardly have failed
to call attention to the powerful second uropods described
by Norman (Shetland Dredging Rep.) under P. hirsuti-
mana had they existed in his species.

Gammaropsis erythrophthalma (Lillje.).

Turbot Hole, Puffin Island, 15 f.; Anglesea coast, 20 f.,
8/6/89; 16 m. of N. of Holyhead, 45 to 50 f.; between
Mersey Bar and Isle of Man, 20 to 30f., 28/8/86; 8m. W.
of Fleshwick Bay, 30 to 33 f. {off Blackcombe; Lambay
Isle}. Length 73 mm.

Gammaropsis nana, Sars.

My specimen differs shghtly from Sars’ description, the
secondary app. of upper ant. being three-jointed (the last
very small) and the dactylus of first gnathopods finely
serrate on the proximal half. The eyes are oval—not
round. Otherwise they agree. 5 miles W. of Dalby,
Isle of Man, 30 f., sandy mud, 8/7/94; 8 miles off Flesh-
wick Bay, 30 to 33 f., 5/6/92. Length of female with
ova 24 mm.

Megamphopus cornutus, Norman.

Podoceropsis intermedia, Stebbing.
Protomedeia longimana, Boeck.

312 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Off Garwick Head, 24/9/92; Little Orme, 28/7/93; 8
miles off Fleshwick Bay, 30 to 33 f, 5/6/92; Menai —
Straits off Beaumaris 10 to 15 f., 17/9/94. None of the
specimens taken so far show the prolongation of the first
epimere of the person described by Norman and from
which this species takes its name. Apparently it only
acquires its full development in northern seas. Colour
transparent whitish sparingly freckled with dull red.
Length 4 mm.

Microprotopus maculatus, Norman.

Colwyn Bay and Little Orme; Puffin Island, 15 fathom ;
Menai Straits; Port Erin Harbour—abundant; tow-net
25 miles 8. EH. of Calf of Man 6/6/92. Colour brown.
Length 35 mm.

Photis longicaudata (Bate).

Puffin Island, 15 fathoms. 8 m. W. of’ Port Mame
tf, mud and 6 im! S.Eot Cali 134 ft camee a and
gravel, 25/4/95. Length 6 mm.

Photts pollex, n. sp. (Pl. XIX., figs. 16—19).

=P. remnhardi, Rep. L., p. 216. Very near P. tenwi-

cornis, Sars, from which it differs only in the second
enathopod of the male. In this the second joint is nearly
half as long as the first and equal to the third. The prop.

has the post. margin prolonged into a tooth or thumb ~
having its base much nearer the carp. than in P. reinhardi

(whence the specific name) the point of which exactly
meets that of the dactylus. The palm, which is longer
than the post. margin, is concave but expands distally
into two tubercles so that this portion of the propodos is
wider than the middle. The female is like that of P.

tenuicornis. Length 2 mm. Colwyn Bay, shore to 23° g

f.; Little Orme; Menai Straits, 5 to 10 f., 24/5/90.
Podoceropsis excavata (Bate) = Nema rimpalmata, :
ep. Leape ait. | ¥:

REVISION OF L.M.B.C. AMPHIPODA. aly

Rhos Bay; Puffin Island; Red Wharf Bay, 20 f., 8/6/89 ;
Port Erin, outside harbour; off Southport. Length 8 mm.

PODOCERIDZ.

Amphithoe rubricata (Montagu) = A. podoceroides, Rath.

=A. luttorina, Bate.

North Coasts of Wales to Holyhead and Isle of Man,
generally under stones between tide marks. Length
13 mm.

Pleonexes ganmaroides (Bate) = Sunamphithoe gam-

marotvdes.

Moelfre Bay, Anglesea in rock pools among Lanuinaria
August, 1889 (F. Archer). Length 8 mm.

Ischyrocerus minutus, Lilljeborg = Podocerus isopus,

Walker, Rep. III. and IV.

Abundant in tidal pools in April, Rhos and Colwyn
Bays; Great Orme; Puffin Island; Menai Straits.
Colour whitish with broad transverse bands of reddish
brown in the females, and numerous small spots of the
same colour in the male. Length of adult male 5 mm.
This species was erroneously united by Boeck with
P. anguipes (Kroyer) which caused me to make a new
species of it. Adult males are rare, and ovigerous females
vary greatly in size. Dredging just below low water
mark in Rhos Bay among stones on May 13, 1893 I took
more individuals of this species than of all the other (22)
species taken together; some of the ovigerous females
measured 33 mm. and others only 23 mm.

Podocerus falcatus (Montagu) (Pl. XIX., fig. 20).

Rhos and Colwyn Bays; Puffin Island; Menai Straits ;
Port Erin Harbour; between Holyhead and Isle of Man,
50 f., 20/7/89, a very large male 10 mm. long—the prop.
of second gnathopod being 4 mm. long—taken in a tow-net
on the bottom. Colour yellow with brown transverse

314 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

bars and spots which keep their colour in spirit. I have
occasionally seen the markings reddish. Length of a
large male from Port Erin Harbour 7 mm.

Podocerus pusillus, Sars,=P. minutus, Sars.

Off Port Erm 6/6/92, 1 male and 2 or 3 females with
ova.. Length 3} mm.

_Podocerus herdmam, Walker (Sixth Ann. Rep. L.M.B.

Com., 1898, p. 37, fig. 138. Brit. Assiepebega.
p. 539). P. odontonyz, Sars, 1894. Bap!

-Off Port Erin from Pecten, &c.; 8 miles W. of Flesh-
wick Bay, 33 f.; 4 miles N. W. of Bradda Head, 21 f.,
29/1/93; Colwyn Bay, tidal pool, 29/1/93. Length 3} mm.
There is no doubt that the above species are identical
as suggested by Prof. Sars, and as P. herdmani was
published first, the name P. odontonyx must lapse.

It is difficult to say whether the two last species (P.
pusillus and P. herdmant) are really distinct from P. fal-
catus. If it be admitted that Amphipoda may become
sexually mature before they have attained their final
moult I think these species can hardly be maintained.
As regards P. pusillus, Sars bases his separation of it
irom whe falcatus on (1) the structure of the post. gnatho-
pods (2) its inferior size and (8) (Oversigt af Norges
Crust., p. 112) its occurrence in deep water, while P. fal--
catus is a littoral or sub-littoral form. As to (1) one ~
need only compare Sars’ figures of the 2 species (and
the accuracy of his drawings 1s remarkable) to see how
slight these differences are; as to (2) the variation in size
of apparently adult individuals is so great that this cannot
be properly used as a specific character; while as to (3)
I have mentioned above the occurrence of a very large
male of P. falcatus at a greater depth than we have
taken P. pusillus. |

The adult male of P. herdmani again closely resembles —

REVISION OF L.M.B.C. AMPHIPODA. 315

the immature male of P. falcatus (see Sars, Pl. 212 p.?
male +). The specimen from tidal pool, Colwyn Bay,
5mm. long has all the appearance of a young male of
P. falcatus but the lower antenne have the last joint of
the peduncle and the first of the flagellum densely clothed
with the plumose sete characteristic of the adult male.
The tooth at the base of the palm of the second gnatho-
pod is more pointed, and that on the dactylus scarcely so
large as in the typical form but hardly two specimens
are exactly alike in these points, especially the latter. As
to the females of all three species, I confess that I am
unable to see any difference between them except in size.

Podocerus variegatus (Leach) (Pl. XIX., fig. 21).

17 miles N. W. of Mersey Bar, 27/9/90; Menai Straits
7 f., 30/5/94. Whitish variegated with red. Length 7mm.
There has been much controversy about this species.
Boeck (Crust. Amph. bor. and arct.) describes Janassa
capillata (Rathke) under the name of J. vartegata, Leach,
while Nebeski (Beitr z. Kenntniss der Amph. der Adria)
unites it with P. falcatus to the female of which it bears
a considerable resemblance in the form of the second
enathopod. It may, however, be at once recognised by
the more robust antenne and the difference in the flagel-
lum of the upper. In P. variegatus the flagellum is four-
jointed, the first jot being nearly twice as long as the
remaining three which are subequal. ‘There is a distinct
secondary appendage which is about +th the length of the
first joint. In P. falcatus the flagellum is seven-jointed,
the first jot rather shorter than the three following;
the secondary appendage about jth the length of the
first joint. In P. variegatus the second gnathopods do
not differ materially in the two sexes except in size, those
of the male proportionally much the larger.

From Janassa capillata (Rathke) this species may be at

316 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

once distinguished by having two secondary teeth below
the curved terminal nail of the third uropods, in which
respect it resembles P. falcatus, and by the distinct,
though small, secondary appendage of the upper antenne,
this in Janassa being so absolutely rudimentary that it is
only visible as a minute tubercle under a very high power—
say ¢ In. obj. All the limbs are proportionally shorter
and stronger than in P. falcatws and the second joint of
the palp of the maxillipedes is more than half the length
of the first jot, while in P. falcatus it is less than half
the length. |
Podocerus ocius, Bate (female)—Della Valle, Fauna des.
Golfes v. Neapel—Gammarini.

From sponge débris, Port Erin. Bate’s figure appear
to have been taken from a female specimen; the male is
figured by Della Valle. It has two pointed teeth or
processes at the base of the palm in the second gnathopods
the proximal one being the longer. The outer ramus of
the third uropods has a curved but blunt nail and no
secondary teeth. Colour brown. Length of adult male
23 mm. )

Podocerus cumbrensis, Stebbing and Robertson (Trans.

Zool. Soc., Vol. XIII, ps 38, Pl. VEY |

Rhos and Colwyn Bays; Puffin Island; Menai Straits.
Colour brown. Length 3 mm. Not uncommon in the
above localities. It has a strong superficial resemblance
to Microprotopus maculatus, with which I have generally
found it associated, and may easily be overlooked as being
the latter species. |

Janassa capillata (Rathke).

Puffin Island; Port Erin, breakwater, 2/8/94, common;
5 m. W. of Dalby, 30 f., 8/7/94. This species may be
distinguished at a glance by the extreme hairiness of the —
antenne ; the flagellum of the upper is three-jointed, the

REVISION OF L.M.B.C. AMPHIPODA. Slt

first joint three times as long as the remaining two. The
first joint of the second gnathopods and two first pairs of
perzeopods are conspicuously yellow. Colour grey with
transverse bars of brown. Length 6 mm.

Hrichthomus abditus (Templeton).

Point of Ayr; Puffin Island, 14f.; Little Orme; Menai
Straits; 16m. N. of Holyhead, 50 f.; Port Erin Harbour,
Length 8 mm.

Erichthonius difformis, M. Kdwards.

Port Erin—electric lhght; Laxey Bay, 4 to 12 f.,
24/9/92, very abundant in tubes on Zostera. Length 6mm.

COROPHIIDZ.

Siphonecetes collettt, Boeck.

Off Garwick Head, 4 to 12 f., 24/9/92; Port Erin Har-
bour; Little Orme. Length 3 mm.

Corophiwm grossipes (Linn).

Mud banks in the estuary of the Dee; tidal ditch in
Rhos Bay; Menai Straits. Occurs in immense numbers
wherever there are mud banks left bare by the tide.
There are hundreds of acres of such banks in the Dee
closely perforated by its burrows. It forms an important
part of the food supply of the various wading birds, and
of fish. Length 7 mm.

Corophiwm crassicorne, Bruzelius.

C. spumcorne—Bate, Brit. Mus. Cat., female.

C’. bonellu, M. Kdw.—Bate and West., Brit. Sess.
Crust., female.

C. crassicorne, Bruz.—Della Valle, Gam. des
Golfes v. Neapel, female.

C’. crassicorne, Bruz.—Bate and West., Brit. Sess.
Crust., male.

Little Orme, 5 to 10 f., 5/10/93. The immature male
has a row of spines on the penult. joint of the lower ant,

‘318 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

as in the female, but this joint terminates in a tooth-like

‘process as in the adult male. Length 3 mm.

Corophium bonelli, Milne Edwards. .
C. crassicorne, Bruz., var., Hoek Tijdschrift Neder-
land. Dierkund Vereen. 4“Deel, 1879, Pl.

VIIL., figs. 9 and 10.

Little Orme; Puffin Island; Menai Straits; Port Erin;
off Clay Head, 18 f. |

Corophium crassicorne of Hoek, l.c. Pl. VIIL., figs. 4
and 5 seems to agree with C. acherusicum (Costa) as des-
cribed by Della Valle. Costa’s description is too vague
for identification. Length 3} mm. |

Unciola crenatipalmata (Bate).

Porthwen and Dulas Bays, 17 f., 8/6/89; off Lianiaelog,
Anglesea, 24/5/90; Penrhos Bay.

For description and synonyms see “‘ Bonnier, Les Amph. -
du Boulonnais.”’ Bull. Scient. de la France, &c., 1889, Pls.
XII. and XIII. This species is closely allied to U. leucopis
(Kroyer) from which it differs in the colour of its eyes
which are dark and very distinct, in the absence of the
nodules on the sides of the pleon, and (according to Sars)
of the transverse ridges of the pereon. This last feature

is not mentioned by Kroyer or Boeck and may possibly

have been produced (as it frequently is) by contraction
after death. Length 5 mm.
U. planipes, Nor.=U. leucopis (Kr.) Bate & West., I., 517.
Red Wharf Bay, 20 f., 8/6/89; Dulas Bay; Little
Orme, 28/7/93; off Southport, 10 to 20 f., June, 91 (1.C.T.)
Length 5} mm.
Colomastiz pusilla, Grube, see Bonnier, l.c. Vol.
OO NE eon MES Zi IOE,
Cratippus tenwipes, Bate.
Haunguia stylipes, Norman (Ann. and Mag. N. H.,
Ser. 4, Vol. ITI., Pl. XXII., figs. 7 and 12),

REVISION OF L.M.B.C. AMPHIPODA. . - 319

Menai Bridge among sponges, 10 f., April and May,
1894. Sponge débris, Port Erin, 1894. Length 3 mm.

CHELURIDA.

Chelura terebrans, Philippi. .
In posts of the old shipping stage, Penmaen, Colwyn
Bay, with Limnoria lignorwm (Rathke). Length 6 mm.

DULICHIIDA.

Dulichia porrecta (Bate).

Rhos and Colwyn Bays shore to 7 f., common; Menai
Bridge, 10 f.; April and May, ’94, common; from Pecten,
7m. W. of Bradda Head, 38 f., 25/4/95. A large female
with ova had the peduncles of the upper ant., the lower
margins of the epimeres, and the pereeopods (except the
last which are variegated with white) dark red. Hggs
white. Younger specimens are only speckled with red,
as are generally the males which are comparatively scarce.
Length 5 mm.

CAPRELLIDZ.

Phtisica marina, Slabber= Proto ventricosa (Miller).

Little Orme; Puffin Island; Menai Straits; Port Erin
Harbour. Length 73 mm.

Protella phasma (Montagu).

Little Orme; Puffin Island; Menai Straits; Port Erin.
Length 16 mm.

Pariambus typicus (Kroyer)=Podalirius typicus.

Throughout Liverpool Bay; Port Erin Harbour. On
Asterias rubens. Length 5 mm.

Caprella linearis (Linné).

Abundant among Alge throughout the district. Length
15 mm.

Caprella acanthifera, Leach,

820 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Moelfre Bay, Anglesea, tidal pools, Aug., 89 (F.A.);

Holyhead harbour, 12/9/89; Port Erin Harbour. Length
11 mm.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
isle:
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.

mno»rkwhnhr rr

a
S.
3.

10.
1.
12.

Ve.
14.
15.
iG:
ie
18.
SE
20.

21.

EXPLANATION OF PLATES.

Puate XVIII.

Figs. 1—11. Nannonyz spinmmanus, n.sp.

. Head.

. First Maxilla.

. Second Maxilla.

. Mandible.

. Maxillipedes.

. First gnathopod of male.
6a.

(on Pl. XIX.) Dactylus enlarged.
First gnathopod of female.

Fourth pleon segment with hind matory of third.
Telson. |

Side view of telson and third uropods of male.
Second and third uropods of female.
Amphilochus melanops, n.sp., Head. .

PLATE XIX,

Ampito melanops, n.sp., first gnathopod.
5 second gnathopod.

es Telson and uropods.
-Photis pollex, n.sp., first gnathopod.
fe second gnathopod.
a third pleon segment.

ne | Telson and third uropods.
Podocerus falcatus (Mont.), flagellum of upper
antenna.
Podocerus varvegatus, Leach, eee of upper
antenna.

o21

A NEW THEORY OF HEARING.
By C. Hersert Horst, Pa.D.

With Plate XX.
[Read December 14th, 1894.]

INTRODUCTION.

KOHLRAUSCH’S experiments described in Wiedemann’s
Annalen (Vol. X.), amongst others, show that two air-
waves separated by a suitable interval of time suffice for
the production of a distinct tone-sensation, the pitch
being determinable by the ear within the limits of error
expressed by the ratio 24:25. Increase of the absolute
number of vibrations with the same time-intervals narrows
the limits of error in determining the pitch by the ear,
till the absolute number reaches 16, beyond which point
a further increase does not appreciably facilitate the
determination.

These results, and especially the first one, disprove,
not only all existing theories of hearing based on assump-
tions as to ‘“‘resonance’”’ or response of sense-hairs, fibres
of basilar membrane, or any parts of the organ ‘‘ tuned ”’
to the same pitch as the sound, but equally disproves all
new resonance-theories of hearing which may ever be
propounded in the future.

Limits of space forbid me here to repeat a description
of the ear and especially of the cochlea. The description
given by Professor Schafer in Quain’s Anatomy (10th
~ edition) is accessible to all, and I have taken that descrip-
tion as correct except in that I have considered both edges
of the tectorial membrane to be attached, the outer edge
being, not free, but attached to the upper surface of the
cells of Hensen, The measurements given (op. cit., vol,

_ 322 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

III., pt. 111, p. 127) are in millimetres and not in micro-
millimetres as there stated. |

The aim of my investigation has been, not to find an
explanation of the sensation of sound, at least not in the
first instance, but merely to lay a solid foundation for
future explanations by considering the physical changes
produced in the ear by the impact of- sound waves upon
the tympanic membrane.

From this point of view it is equally important to follow
out to the end the whole chain of effects in the case of
every part, however little may seem the probability of
finding in them a solution of the physiological problem
of hearing: the problem to be attacked is indeed in the
first instance not a physiological one at all, but one in
pure physics, and as such it must be treated; leaving
altogether out of account any and every physiological
consequence which may follow, and carefully avoiding all
those suggestive analogies with musical instruments
which have led previous investigators into erroneous

assumptions as to “‘functions”’ of one part or another.

Each part must be considered, in the first instance, as an
inert body whose properties are to be determined only
by observation and not assumed beforehand on the
strength of any preconceived notion as to the way in
which they ‘‘ must’ act in order to produce certain effects.

For this reason the first portion of the theory is
addressed to physicists rather than to physiologists and
to the latter only is so far as they are physicists as well
as physiologists.

Tum PHYSICAL EFFECTS OR CHANGES PRODUCED IN THE — ;
MIDDLE EAR BY THE Impact oF SoUND WAVES
ON THE TYMPANIC MEMBRANE.

The movement of the membrane, due to changes of

A NEW THEORY OF HEARING. aa

pressure on its outer surface while the pressure on its
inner surface remains nearly constant, produces effects
upon the ossicles of the middle ear. To determine
whether the ossicles move as a whole or as some have
maintained they ‘‘ transmit the sound molecularly”’ it is
necessary to make a rough estimate of the velocity of
sound in them and to consider what is the length of a
sound wave when transversing the series of ossicles.

To determine this accurately is not easy but it is quite
unnecessary: an error of even 10000 per cent. is of no
consequence. The lowest audible tone 1s not the same
for all hearers, but few if any can hear as a tone any
sound due to less than 10 vibrations a second. A pure
tone of 40000 vibrations is audible to some and the pro-
duction of beats and beat-tones by simultaneous action of
two sets of waves some of which in one series follow
others in the other series at smaller intervals of time than
this has been demonstrated by Koenig and others. It 1s
only necessary therefore to consider whether the time
required for transmission of a sound wave through these
ossicles lies between ssganth second and {5th second, or
above or below those limits. The mere consideration of
the question even without answering it, suffices to show
that the distinction drawn between the two modes of
transmission is a distinction in words rather than in
phenomena. If the wave length in the ossicles is much
smaller than the length of the chain it would be said that
the transmission is “‘molecular;” if it is much longer
than the chain it would be said that it is ‘‘molar.’”’ The
nature of the transmission is, however, identical in the
two cases and the distinction is purely verbal and useless.
The transmission of energy along a pulled bell-wire is of
the same nature as the transmission of a sound wave:
the energy is indeed transmitted as a true sound wave of

324 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

enormously long period and enormously great amplitude :
the wave length being enormously great as compared with —
the length of the wire. It is only necessary therefore to
point out here that the movements of the base of the
stapes follow those of the tympanic membrane but lag
slightly behind them: how much they lag probably
depends on the wave-length and on the intensity of the
sound: the lag is probably less with intense sounds than
with fainter ones. It probably amounts to less than a
wave-period with low tones and to more than a wave-
period with high tones. With very violent waves of long |
period the elasticity of the displaced membrane, the
inertia of the ossicles, and the tension of the muscles and —
ligaments attached to the ossicles as well as the nature of _

the joint (saddle-shaped) between the malleus andincuswill

come into play and reduce the amplitude of displacement —
to a very small fraction of what it would otherwise be.
The scale of relative amplitudes of displacements of them
fenestra ovalis will therefore be very much smaller than that _
of the displacements of the tympanic membrane. Veryfaint —
sounds, in other words, will produce a relatively far —
greater effect on the ovalis than very loud ones: nearly |
the whole energy of high tones and of faint sounds will —
be transmitted to the ovalis, but only a very small fraction |
of the energy of more violent waves and especially of waves —
of great amplitude in the absolute sense (e.g. explosion-
waves). Thisis a highly important protective arrangement —
and is analogous to the protective action of the iris in
the eye.

THE CHANGES IN THE INNER Har.

The whole inner ear may be rightly regarded as a rigid

box with only two yielding areas in its walls and filled —

with an incompressible fluid. The two yielding areas are
the fenestrae rotunda et ovalis,

A NEW THEORY OF HEARING. 325

The errors involved in so regarding it have been roughly
estimated and their effect on the conclusions which follow
is ml. The chief of them is involved in the assumption
as to the incompressibility of the fluids (perilymph and
endolymph). These fluids are almost identical in physical
properties with water and if we consider the modifications
of the following account which would have to be made in
the case of water we shall see that the error is one which
does not affect the theory as a whole.

Water is compressed to the extent of rather less than
sooooth part of its volume by an increase of pressure equal
to one atmosphere. No such pressure as this is ever
brought to bear on the fluids in the ear; if it were the
transmission of the effects, 2.e., of the movements, would
be accelerated. As I shall give a justification for the
assumption that this transmission is w/initely rapid
nothing tending to increase that velocity can possibly
affect our conclusions except in the direction of increased
accuracy.

The velocity of sound in water is almost one mile per
second. The distance between the fenestre ovalis and
rotunda is about 4 inch. ‘The time required for trans-
mission of a pressure wave (‘‘sound wave’’) from one
fenestra to the other is less than zopapath of a second,
2.€., less than 75th part of the time-interval between two
successive waves of the highest audible note. I shall
regard this interval as nz/, and shall regard the movements
of the two fenestre as simultaneous, which is equivalent
to regarding the velocity of sound in the fluids as infinite
and the fluids themselves as incompressible. It will be
seen later on that the error involved in this assumption
does not, even in the most minute degree, affect the
soundness of the conclusions.

The assumptions being justified I will take a series of
problems in the following order :—

326 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

1. The changes produced within the cochlea by one
inward movement of the stapes followed by its
immediate return to its position of rest.

2. The changes produced by two such double move-

| ments. . .

3. The changes produced by a long series of such
movements at equal intervals of time—as by the
impact of the sound-waves due to the sounding
of a constant pure tone.

4. 'T'he changes produced by the simultaneous action
of two such series of waves due to sounding two
notes in harmony.

5. The corresponding problem in the case of discordant
tones.

Problem I. Changes produced within the cochlea by
one double movement of the stapes (in, then out).

The stapes, and of course the fenestra ovalis in which
it is embedded, moving inwards, the rotunda must on the
foregoing assumptions move outwards simultaneously. ;
This involves movement of the fluids between. This
movement is interfered with by the membranes lying
between the two scale, upper and lower (2.e., scale
vestibuli et tympani), of the walls of which they form
parts, and the displacement of these membranes together
with the subsequent effects of those displacements presents _
the fundamental and most important problem we shall —
have to consider. |

For convenience the cochlea will be regarded as a spiral —
with its axis vertical and the apex at the top: the basal
region is then the region where the fenestre are placed :
the three scale, tympani, media and vestibult become —
the lower, middle and upper canals and the basilar and
Reissnerian membranes become the lower and upper
membranes respectively.. Movement. along a canal will —

A NEW THEORY OF HEARING. 327

be spoken of as forwards or backwards according as it is
towards the apical or towards the basal region.

First as to the upper membrane. The fenestra ovalis
is placed at the junction of vestibule and cochlea.
Pressure applied to it affects the fluids within and around
the membranous vestibule equally, as also around and
within the canalis rewwens. Any movement of the upper
membrane near the junction of the walls of the canalis
reuniens with it will therefore be due to friction only and
not to difference of pressure on its two surfaces. This
portion of the upper membrane is moreover loose and
flabby and the movements which can only be slight will
involve no stretching. The membrane is moreover light,
broad, thin, soft and relatively inelastic (on account of its
looseness at this point) and it will move therefore freely
as part and parcel of the fluids in which it is immersed,
such elasticity as it possesses not being brought into play.
The effects of a wave of this membrane if one should be
produced would be identical with, but enormously smaller
than, the basilar wave to be considered directly. For the
present therefore we may without error ignore the
existence of the upper or Reissnerian membrane, and look
upon the basal portion of the cochlea as consisting of only
two chambers or canals separated from each other by the
basilar or lower membrane.

The lower membrane is very different. It is exceedingly
narrow, its basal end may be said to end almost in a
point: it is not only exceedingly stiff by virtue of its
transverse fibres, but its elasticity is enhanced by the
transverse tension under which it is stretched between
the two bony ridges supporting it. A very slight dis-
placement will therefore instantly call its elasticity into
play, and the movement towards the lower canal affecting
first its most basal portion will rapidly extend along the

a a PEE

See i

328 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

membrane during the whole period of positive acceleration
of the inward movement of the stapes, tension being
sreatest at the basal end. During the second portion of
the advance of the stapes (with a negative acceleration)
this first portion of the lower membrane will commence

to return to its position of rest, a longitudinal movement

of fluid in its neighbourhood being involved therein. This
movement of fluids will be forwards in the upper canal and
backwards in the lower canal but will extend through only a
small distance, and will affect chiefly the portion. of fluid
very near to the surfaces of the membrane, but not quite in
contact with them. The reverse or the return movement
of the stapes will produce a similar effect in the opposite
direction, the first portion of the return movement of the
stapes augmenting the fluid movements of the second part
of the advance. If we now consider the condition of the ~
parts near the base at the moment when the return
movement of the stapes is just completed we shall find
that they are such as are necessarily transmitted in the —
form of a wave, though a wave very different from either
a sound wave or a wave in water or any wave which has
hitherto, so far as I know, been considered by phy
or mathematicians.

Let ABCDH, in fig. 1 (Pl. XX-); represent the middle
line of the displaced membrane, A being the basal end of
it. The portion AB is moving downwards to its position
of rest, the fluid displaced by it below being driven for-
wards (v.e., away from the basal end of the canal and
towards the apex); the fluid above moving downwards
with the membrane and somewhat backwards, its place
being filled by fluid from beyond B.

The portion BC is moving upwards, being carried by
the moving fluid below which is driven forwards by the
downward movement of AB. The portion CD is returning

A NEW THEORY OF HEARING. 329

by virtue of its own elasticity towards its position of rest,
to be immediately carried beyond that position by the
movement of the fluid, below it; while it is itself forcing
the upper fluid forwards thus depressing the portion DE,
The small arrows in fig. 1 show the directions in which each
portion of the membrane is moving at the moment, viz:—
AB and DE are moving downwards the points A, B, D,
and K being momentarily at rest, A and E in equilibrium
Band D at maximum tension. BC is moving upwards
with increasing tension and diminishing velocity; CD
upwards with diminishing tension and increasing velocity.
At C is the point of no tension and maximum velocity.
The large arrow shows the direction in which the wave
will travel forwards. |

Fig. 2 shows in a generalised and simplified way the
movements of the fluids. In the upper canal the fluid is
moving from the central wave-region outwards in both
directions, 2.e., forwards from N to M and backwards
from N to O. In the lower canal the movements are the
opposite of these, 7.e., ato the central region N! from
M' in front and from O! behind.

The wnovements of the fluids in each region are first
arrested by the elasticity of the membrane and then
reversed: that is, the kinetic energy of the moving fluid
is transformed (so to speak) into potential energy of
stretched membrane, and then again into kinetic energy
of moving fluid to be similarly transformed in a portion
of the membrane further along the canal.

It may be asked why the disturbance is transmitted
along instead of across the canals; mcompressible fluids
in rigid vessels can obviously however only move in such
directions as the yielding areas of the walls allow: if fluid
is forced forwards in the upper chamber a corresponding
and equal movement of the fluid in the lower canal must

830 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

occur in the opposite direction. The upper and lower
canals communicate freely at the apex of the spiral and if
the displacement of the stapes were infinitely slow the
flow would be a steady one alike in all parts of each
canal, 2.e., during advance of the stapes it would flow
forwards (v.e., towards the apex) in the upper and back-
wards in the lower canal. Or if the basilar membrane
were perfectly rigid the same result would follow.

With a rapid movement of the stapes and with a
yielding though highly elastic basilar membrane the case
ig very different: the inertia of the whole column of fluids
in both canals prevents their sudden movement and only
the basal portion is moved first, the rest being only
subsequently set in movement by the elasticity of the
membrane called into action by its displacement.

The figures 1 and 2 and the constant reference to the
elasticity of the displaced membrane are liable to suggest
a longitudinal tension comparable with that of a vibrating
string. There is, however, practically no such longitudinal
tension. The stretched fibres he transversely in the
membrane, and each stretched portion of the membrane
acts on adjoining parts of the membrane only through the
intermediation of the incompressible fluids in which the
whole membrane is immersed, fluids which are prevented
from moving transversely by the rigid walls of the canals.

We have here the conditions necessary for transmission -
of a wave-like disturbance: we have indeed conditions
which render such a transmission inevitable. The con-
ditions at C for instance are precisely those which at an
earlier moment obtained at B and will in a moment later
hold at D. The whole of the conditions are such as must
give rise to like conditions in an adjoining portion of the —
membrane further forwards. That is the disturbance will
of necessity give rise to a like disturbance in every

A NEW THEORY OF HEARING. Sod,

successive region of the membrane and adjoining fluids
throughout the whole length of the spiral.

This disturbance I will call a wave, in spite of its
difference from any other known kind of wave in the
following respect, viz., the potential energy is that of fibres
stretched at right angles to the direction of transmission,
and the kinetic energy is that of moving fluids; while in
other waves the kinetic and potential energy consist in
movement and stress of one and the same medium. ‘The
inertia of the fibres in the basilar is here insignificant
while in the vibrating string of a fiddie or of a cord thrown
into waves, or of a solid, liquid or gas transmitting a
sound wave or of the water in an oceanic wave the inertia
is all-important. Expressed in another way the whole of
the kinetic energy of this wave is in the fluids; the whole
of its potential energy isin the membrane. The kinetic
and the potential are equal, and the whole energy of the
wave remains constant throughout its transmission, save for
a portion, which must be very small, and which is converted
by internal friction into heat or other disturbances.

Two important questions now arise which are intimately
connected with each other; they are :—

(a) What is the extent or amplitude of na Mlereemne a
of each part of the membrane? and (b) what is the velocity
of transmission of the wave? Neither question admits
of a very definite answer but it is important to give a
partial answer to both.

The spiral canals of the cochlea are not of uniform
width throughout, but are much narrower in the apical
than in the basal region. Suppose for a moment, what
is not true, that the basilar membrane is of uniform
width ; then the wave in passing from the wider to the
narrower part of the canal would gain in amplitude: for
at each successive level the yolume of fluid to.be moved

-
; 7%

332 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

would diminish and its inertia decreasing the expenditure
of like energy upon it would give it a greater velocity and
therefore a greater displacement—an effect comparable
with the ‘‘ bore” in the Severn and similar rivers.

The basilar, however, is not uniform in breadth: it is
about twelve times as broad in the apical region as in the
basal: that is the wave in passing forwards towards the
apex comes into successively narrower and narrower
portions of the canal and acts upon successively broader
and broader areas of the basilar membrane. The effect
of this is to exaggerate the effect of the narrowing canal
already described. In order that the same amount of
energy may be stored in potential form—to speak figura-
tively—in two fibres which are otherwise alike but of
different lengths the transverse displacement of the longer
one must be much greater than that of the shorter one.
The wave in passing forwards towards the apex will thus
rapidly gain in amplitude—and the importance of this
fact from a physiological point of view will be seen later.

What changes in velocity result from this change in
width of the channels and in breadth of the membrane
as the wave passes on it is less easy to say. The two
influences—narrowing of canal and consequent decrease of
mass per unit length to be moved, and broadening of
membrane and consequent diminution of elastic pressure
- per unit displacement—appear to counteract each other,
but whether the two effects balance each other or one
over-balances the other is a question which I have as yet
only guessed at on physiological and not on physical
erounds. The guess which will be justified later is that
the velocity of the wave and consequently its length (in
space, not in time) will be augmented.

As to the absolute velocity at any part of its course—
as distinct from its relative velocities in different parts of

A NEW THEORY OF HEARING. aa0

its course—we are again driven by the difficulties of the
physical problem to base an estimate on physiological
considerations: what that estimate is will appear later.

The cochlear canals, however, are not only tapered but
but also spirally coiled; and the basilar membrane is at
the apex continued into the Reissnerian which runs from
this point to the base of the spiral narrowing very gradually.
When the wave reaches the apex of the cochlea it will pass
on to the upper (Reissnerian) membrane, for at this point
the lower canal is continuous through the helicotrema
with the upper (scala vestibuli) ; and the middle canal ends
blindly in the slightly dilated “‘lagena,”’ the wall of which
is formed by the two membranes (upper and lower) which
are here continuous with each other.

The wave of the basilar will thus, starting from the
base of the cochlea close to the fenestra rotunda, run
forwards to the apex, round the wall of the lagena and
down by the Reissnerian membrane to the base of the
cochlea.

The elasticity of the Reissnerian membrane being much
less than that of the basilar, it appears certain that the
displacement at each point will be greater than that of the
corresponding portion of the basilar, and further the loss
of energy of the wave will be greater for the movements
involved being more extensive will involve more internal
friction. The smaller waves will hence probably be almost
lost before reaching the base, and the larger ones will,
apparently, die out at the base in the form of small
vortices or eddies of the perilymph gradually coming to
rest by internal friction of the somewhat viscid perilymph.
This is not the place to discuss the far greater effect of
viscidity of a fluid in restraining vortical movements than
in restraining wave-like movements which involve only a
temporary and relatively small internal deformation (or

304 TRANSACTIONS LIVERPOOL BIOLOGICAL ‘SOCIETY.

relative displacement of parts) of the fluid. It is sufficient’
to state that such is the effect of viscidity, and that it will
rapidly arrest such vortices as would be formed when the
Reissnerian wave reaches the base of the cochlea.

The effect of the spiral coiling of the cochlear canals
will be to concentrate the disturbance in the outer part of
each turn of the spiral. The large, longer waves will thus
involve a disturbance of a larger mass of fluid at each
level, while the smaller ones will involve movements more
narrowly restricted to the outer region of the canals that
is, to the region where the membranes are placed.

The effects of these waves on the organ of Corti have
now to be considered, and the effect of the passage of a
single such wave as has been described will be seen to be
nil, or rather that it will consist only in a bodily movement
of the whole organ in each region first down then up and
then down again, with little or no relative movement of
the parts of the organ among themselves.

The upper surface of the hair-cells is covered by the
comparatively stiff and tough reticular membrane connected
with the phalangar processes of the rods of Corti, and the
whole covering so formed is rigidly connected with the
basilar membrane by the stout rods of Corti and the less
stout, but still stiff “cuticular rods” of the cells of
Deiters. This skeleton, as it may properly be called,
insures the movement of the basilar membrane and the
whole mass of hair-cells and cells of Deiters as one piece
without deformation or relative movement of its parts.
The whole organ in fact in each transverse area may be
regarded as part of the basilar membrane, moving up and
down but retaining its form.

The tectorial membrane, however, hes upon this and is
connected with it only at its edges. It is often described
as being free at its outer edge, but this edge always

A NEW THEORY OF HEARING. — 335

presents the appearance of being torn and jagged and
portions of it may sometimes be seen attached to the
upper surface of the cells of Hensen: a portion so attached
was figured by Retzius and is shown in the figure repro-
duced from his work in such text-books as Foster’s
Physiology (fifth edition, p. 1851). It is highly probable
therefore that this membrane is attached at both edges
throughout the entire length, and that its frequent
separation is due to shrinkage under the action of reagents.
A similar shrinkage of the upper surface of their hair-cells
is prevented by the stiff unyielding reticular membrane,
and the tearing of the tectoria at its thinnest part when
exposed to reagents which cause shrinkage is precisely
what might be expected—it might even be said to be the
inevitable consequence.

The tectoria when stretched in its natural condition
between the cells of Hensen and the lip of the spiral
lamina would rest just over the tips of the hairs of the
hair-cells. Between the tectoria and the reticular mem-
brane is then a thin layer of fluid which is somewhat
viscid and in which the stiff hairs project. The thickness
of this layer is about ‘004 mm. and in consequence of its
viscidity the friction involved in its movement would be
relatively great. The fluid in the wide scala media above
the tectoria 1s, however, much more free to move and the
tectoria would therefore move with the organ of Corti
almost as if it were rigidly bound toit. The passage of the
basilar wave would thus simply move the whole organ of
Corti, together with the tectorial membrane up and down
without disturbing the relative position of one to the other.

This purely physical conclusion accords well with the
physiological observation that a single aerial wave 1m-
pinging upon the ear does not suffice to produce a
sensation of sound.

836 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

Problem II. Changes produced within the cochlea by
two double movements of the stapes (in then out.)

If the first wave has already traversed the whole length
of the basilar first and then of the Reissnerian, and finally
died out before the second wave is started, the second
will obviously pursue the same course as the first and
produce only the same effects. This consideration enables
us to roughly estimate the average velocity of the waves
along the membranes; for if the time-interval between
the two waves be progressively diminished in successive
experiments, a minimum period of no physiological effect
will ultimately be reached, and as soon as this is passed a
sensation of sound is produced. That minimum time-
interval of no physiological effect thus corresponds to the
time occupied by a single wave in completing its whole
course up the basilar and down the Reissnerian mem-
branes. This time-interval differs with different persons,
normally ranging between #th and 4,th of a second.

When the time-interval is less than this the second
wave will be started on its journey up the basilar mem-
brane while the first wave is still somewhere on one or
other of the membranes; and before the second reaches
the helicotrema it must of necessity pass some point on
the basilar membrane at the same moment as the first
wave passes a point opposite to this on the Reissnerian.
Two points have to be considered in connection with this
passing of the waves; (1) the point at which the passing
will occur, and (2) the effects within the organ of Corti
which will be produced thereby.

(1) The velocity of transmission of a wave at any given
point of either membrane being constant (as it must be
very nearly if the waves are similar to each other), it
follows that the distance from the position of the second
wave measured along the basilar through the helicotrema

A NEW THEORY OF HEARING. 307

and then down the Reissnerian to the position of the
first wave, is the distance which has been travelled by
the first wave since it was in the position now occupied
by the second ; 7.e. it 1s the distance travelled by the first
wave in the interval of time between the first and the
second. This distance thus depends entirely upon the
time-interval between the two waves. As the time-
interval between the two determines the pitch of the
tone-sensation produced by them (as shown by Kohl-
rausch’s experiment referred to above) it is correct to say
that if a note be sounded consisting of two vibrations only
(in the absolute sense) then the point in the organ of
Corti where the two cochlear waves pass each other will
depend wholly upon the pitch of the note. In other
words, there is in the organ of Corti for each pitch a
definite point where this passing will occur. We shall
later see that though there is only one such point when
only two vibrations are produced, there may be a larger
number when more than two are produced.

(2) The effect on the Organ of Corti and associated
structures produced by the simultaneous action of these
two waves may be most easily grasped by the mind by
consideration of a transverse section of the middle canal
at this level. First both upper and lower membranes are
motionless and the fluids are at rest. Then the two are
simultaneously drawn or forced apart, and coming to rest
for a moment then move suddenly towards each other
driving the fluid out of this region in both directions.
A more accurate idea may be obtained from consideration
of a longitudinal section through the mid-line of the two
membranes in this region.

Figures 3, 4 and 5 represent three phases of the passing
of two such waves. They will be more easily understood
if compared with figures land 2. The large arrows above

- 338 TRANSACTIONS LIVERPOOL BIOLOGICAL’ SOCIETY.

and below represent the direction in which the first and
second waves are travelling along the upper and lower
membranes respectively. The smaller arrows of figures
1 and 2 are omitted. The line XY which is the same in
all three figures marks the level at which the two waves
pass each other in corresponding (not “like’’) phases.
The first wave, in passing the helicotrema has of course
been turned over so that its first phase is one of woward
displacement, while that of the second wave which has
not been reversed 1s one of downward displacement. Let
points in the waves be called ABCD and E as in fig. 1,
and phases M, N and O as in fig. 2—it is more convenient
to divide the wave into these three phases than in the more
usual way. In fig. 3 at the level XY both waves have
just reached the end of the first phase (M), 7.e., the point
D of each wave is just passing the hne XY. This means _
that fluid has just been forced into region of XY from
both before and behind, and in the neighbourhoods of both
upper and lower membranes. ‘The two points B of upper
and lower membranes are now separated to the utmost
extent and by virtue of their elasticity have just brought
the fluid in contact with them to rest. From this moment
onwards the two membranes by virtue of that same
elasticity will approach each other, with increasing velocity
till at the level XY the stage shown in fig. 4 is reached.
The points C (fig. 1) of the two waves are now passing
each other at the level XY. At this moment the two
membranes are approaching each other with maximum
velocity, being forced towards each other by the fluids in
the upper and lower canals, and the fluid between the
two membranes, that is in the middle canal (scala media,

canalis cochlez) is being forced out from this region, now
with maximum velocity, both forwards and backwards.
A moment later the stage shown in fig. 5 is reached.

A NEW THEORY OF HEARING, - - 3389

The points B (fig. 1) are now passing each other at the
level XY. The two membranes at this level have just
come to rest by virtue of their own elasticity, and the
fluid will now immediately commence to flow in both
directions (forwards and backwards) into the region of XY.

We must now consider how these changes will affect
the organ of Corti.

Let fig. 6, represent a transverse section across the
middle canal of the cochlea in the plane of the line XY of
figures 3, 4 and 5, and let us consider the changes Just
described from a new point of view.

The condition represented in fig. 3, was brought about
by inrush of fluid into the region of XY in both directions
sumultaneously. From this stage onwards the elasticity
of the membranes now stretched to their maximum
presses on the fluid between them, forcing it out in both
directions (forwards and backwards) with increasing
velocity till the stage represented in fig. 41s reached, when
the membranes are at their normal position of rest but
are now advancing towards each other with maximum
velocity. During this first half of phase N (fig. 2) the
basilar membrane has been pressing upwards upon the
fluid which was simultaneously forced down upon it by
the Reissnerian membrane, with the result that not only
the fluid above the tectorial membrane but the whole of
the fluids between the basilar and the Reissnerian will
have been set in movement backwards and forwards away
from this region. ‘The whole phase only lasts a moment,
but during that moment the fluid between the tectorial
membrane (T in fig. 6) and the organ of Corti (C in
fig. 6) will have been to some extent forced out, and by
way of the spiral sulcus to adjoining regions of the cochlea.
In other words the tectoria will have been suddenly banged
down upon the hairs which extend vertically between it

840 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and the top of the organ of Corti. These hairs are stiff
and the ultimate fibrils of the auditory nerve terminate at
their bases. This means that at the particular point in
the length of the organ of Corti where the two waves pass
in corresponding phases the ends of the stiff hairs of
the hair-cells of the organ of Corti are suddenly thrust
down upon the nerve-ends of the auditory nerve-fibres.
Physiologists will understand what result will follow.

Taking a general review of this second problem, the
result may be stated thus: When two sownd waves im-
pinge upon the tympanic membrane at a surtable interval
of time, one definite region of the organ of Corti will be
stimulated by the thrusting of the hairs of its hair-cells
upon the nerve-ends ; and the region to be so stumulated.
will be deternuned wholly by the interval of tume between
the two sound waves, that is, upon the pitch of the note
sounded.

Problem III. The changes produced within the cochlea
by a long series of in and outward movements of the stapes,
such as would be brought about by the wmpact upon the
tympanic membrane of the sound waves produced by the
sounding of a constant pure tone.

~ Tf the interval between successive vibrations be of such

length that two and only two cochlear waves were passing
along the membranes of the cochlea at any one moment,
the problem is identical with the previous one. When the
intervals are much shorter the problem becomes more
complex ; and certain details have to be considered which
were for the sake of simplicity omitted from consideration
in the previous problem. In the first place there will no
longer be one place and only one where the cochlear
waves on the two membranes will pass each other in
corresponding phases. They will pass at many such places

A NEW THEORY OF HEARING. 341

and we have to consider whether the results of such
passing as set forth in the previous problem will be brought
about at all these points or only at some of them; and if
only at some, at how many, and at which.

It has already been shown that the effect of the
mechanism of the middle ear is to “level down” the
amplitudes of the displacements brought about by notes
of very different intensities, very intense (v.e. loud) sounds
being prevented by this mechanism from producing a
very much larger displacement of the fenestra ovalis than
is produced by a much less intense sound. Pure tones
of the same pitch but of enormously different intensity.
are by this mechanism caused to give rise to displace-
ments of the ovalis which though greater for intense
sounds than for weak ones, are far more nearly equal than
are the displacements of the tympanic membrane. We
will first suppose that sounds are under consideration
ranging between the very faintest audible tone on one
hand, and a tone of such intensity as will cause double
the amplitude of displacement of the basilar produced by
that faintest audible tone of the same pitch. It must not
be supposed that this means that the second (upper) limit
of intensity represents an intensity of sound twice as
ereat as the lower limit. To suppose this would be to
ignore what has been said about the mechanism of the
middle ear. A two-fold increase of amplitude of displace-
ment of the basilar will correspond to an increase of
intensity of the exciting sound greater than two-fold.

The effect upon the amplitude of displacement of the
basilar arising from the narrowing of the canals towards
the apex of the spiral and the cooperating and intensifying
effect of the widening out of the basilar itself towards the
apex have already been referred to, The net result of

342. TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

these two conditions is that the amplitude of displacement
becomes rapidly greater as the wave passes forwards along
the basilar while it diminishes as the wave returns along
the Reissnerian. For any given series of waves therefore
due to the continuous sounding of a note of constant pitch,
and constant intensity, the maximum effect upon the organ
of Corti will be produced at the uppermost only of the
many levels at which waves pass each other on the two
membranes in corresponding phases. If, therefore, the
damping effect of the mechanism of the middle ear be of
proper extent then the effect upon the organ of Corti
already described will occur at this one uppermost only.
That is, though waves pass at many points it is only at
the uppermost of them that the disturbance will be
sufficiently great to bring the tectorial membrane down
upon the hairs of the hair-cells with sufficient force to
stimulate the nerve-ends.

Equal intensity of sound in the case of two notes of
different pitch, does not correspond with equal amplitudes
of displacement of the air or of the tympanic membrane.
Nor does it correspond with equal displacements of the
basilar membrane. When two sounds—both pure tones—
of different pitch are said to be of equal intensity, that
means that the energy traversing unit area of a plane at
right angles to the direction of transmission in unit time
is equal in the two cases. This being the case two tones
of equal intensity but different pitch involve displacements
proportional to the wave-lengths. A very low note will
therefore involve a far larger displacement than a much
higher note of equal intensity. Still supposing the limits
of intensity above laid down, suppose two notes are
sounded 2% succession, a high one and a low one: and
suppose the intensity of the-two to be at first equal: then
the higher note will produce a stimulation at one point of

fa
tae

A NEW THEORY OF HEARING. 343.

the organ of Corti and the lower at another point. The
cochlear waves due to the higher note besides passing
higher up may pass at the exact point where those of the
lower note do, but the displacement is too small to bring
the tectoria down upon the sensory hairs at that point
although the amplification of displacement as the wave
passes forwards suffices to produce this effect further on
in the cochlea. The lower note on the other hand
produces in the upper part of the cochlea a larger dis-
placement of the membranes than does the higher tone,
but it produces no stimulus because there is no passing of
waves in corresponding phases in these upper regions the
waves being too long to so pass here; that 1s one wave
goes up to the helicotrema and travels backwards along
the upper (Reissnerian) membrane out of the upper
regions of the cochlea altogether before it meets the next
following wave on its way up the basilar.

Within these or some such limits, therefore, each pure
tone will produce a stimulation of the nerve-ends in one
region of the organ of Corti and in one region only: and
the region stimulated will depend solely upon the pitch
of that tone.

Now suppose the intensity to mecrease beyond the
supposed limit.

We shall now have to give names to every point in the
length of the organ of Corti. The number of points is
infinite and there is no difficulty in providing them with
an infinite number of easily understood names or numbers.
Let 100 be the name or symbol of that point in the length
of the organ where one wave in its backward or return
journey along the Reissnerian membrane passes a wave
on the basilar separated from the first by an interval of
time equal to zj5th part of a second. Then within
previous limits of intensity, 100 indicates the point in the

344 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

length of the organ of Corti where a stimulation is
produced by a tone due to 100 vibrations per second. |
Similarly let a stand for the uppermost point of passing
of waves produced by a tone of a vibrations per second.

A tone a will thus produce cochlear waves passing at
points a,4a,4a,+a, etc. Within prescribed limits stimu-
lation occurs at the point a only: at $ @ the amplitude —
of displacement is too small to produce stimulation. |
- A tone 4a of equal intensity is, however, sufficient to
produce a stimulation at the point 4 a: therefore if the |
intensity of the tone a be so far increased as to produce a
cochlear wave of more than double its previous amplitude
(which means manifold more than double intensity), it also
will be able to stimulate the organ of Corti at the point
+ a, but in the first instance the sensation due to the
stimulation at @ will have been so coon intensified
as to overpower this stimulation at +a, and the weak
second tone + @ will be heard either very faintly or not
at all. |
- [have found by experiment, however, that if a tone a —
be sounded loudly for five or ten minutes and then
increased gradually in intensity, the note $a, an octave
lower gradually becomes audible. It is therefore possible,
by gradually deafening oneself to atone and then increasing
its intensity, to produce a sensation of a tone an octave
lower.

The result arrived at in considering this third problem
is thus as follows :—

A continuous constant tone within moderate limits of
intensity will produce stimulation of one region of the organ
of Corti and of one only: and increase of intensity beyond
these limits produces a stimulation of a region correspond-
ing to a tone an octave lower; but this second stimulation
is so feeble as to be difficult or impossible of recognition,

A NEW THEORY OF HEARING. 845

being overpowered by the primary stimulation, until the
first region is deprived of its sensitiveness by fatigue. It
is theoretically possible to fatigue even the second point
-and then hear a spurious tone whose wave-length 1s three-
times that of the actual note sounded. Experimentally I
have failed to produce this effect and this is probably due
to the enormous damping effect of the mechanism of the
middle ear upon sounds of such deafening intensity.

Problem IV. Lffects within the cochlea due to simul-
taneous sounding of two continuous mutually harmonious

pure tones.

The effects arising in various parts of the cochlea when
two series of periodic movements of the tympanic mem-
brane occur simultaneously will differ widely in the cases
when the “‘period”’ of one series bears different relations
to the period of the other. We will consider first the
case of two series the periods of which bear a simple
relation one to the other.

We have already seen that a tone due to @ vibrations
per second may produce a stimulation in the organ of
Corti at points a, 4 a, 4 a, etc., but that the primary
stimulus at @ will be by far the most intense. If while
this tone 1s being sounded, the tone % or % be also sounded,
even though so faintly as to be in itself inaudible, its cochlear
waves may, so far as they happen to coincide with certain
waves of the series a so increase the disturbance at
¢ or % (as the case may be) as to render this tone distinctly
audible. Similarly if the tone a@ (t.e., a vibrations per
second) be sounded too faintly to be audible, and the tone
2a simultaneously sounded so faintly that it would be
inaudible if sounded alone, each tone may so cooperate
with the other as to render both audible, but not quite in
the same way in the case of the two,

if they be of unequal length they cannot coincide in all

346 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

That one wave may intensify the disturbance in the
relative positions of tectorial membrane and sensory hairs
produced by another wave, the two must coincide with
each other in phase in some part of the wave. Of course

parts. It would take us too far to consider the question
as to whether two series of sound-waves of simply-related
periods do tend to arrange themselves in a definite way
with relation to each other or not, and if so what that
relative position is. Fortunately we need not consider it,
for a study of the effects of combination of the waves due
to two tones in harmony with each other by simple
eraphic methods shows that, however, they be combined
the combination will lead either to prolongation of the
period of advance of the two membranes towards each
other at the level of passing of the waves of either series
or to acceleration of the advance. Hither of these effects
will lead alike to the augmentation of the relative move-
ment of tectoria and sense-hairs, at that point. Sounding
of two tones in perfect harmony with each other will
thus produce a slightly greater effect at each of the two
passing-points in the cochlea than the sounding of either
of the two alone.

An effect differing from this, so ie as the sensation is
concerned, may be produced at one or more other points.
Let the two notes be c' and g’ (7.e., the middle ¢ of the
pianoforte and its fifth) and let the musical interval
between them be a trwe fifth, z.e., let the ratio of the
numbers of their vibrations be as 2:3.

Then the points of the organ of Corti stimulated in the
way described will be the points called Gn accordance
with the simple nomenclature I have suggested) ¢’ and
g' or c' and 3¢ The series of waves c’ pass in corres-

2

A NEW THEORY OF HEARING. 347

ponding phase at c’, > = = etc. The second series
pass in the same way at g’, By be = etc. These

points expressed in the simplified form are c', c, F, C and.
9,9, ¢, G, (notation of Helmholtz, Lord Rayleigh and
others) and it will be seen that the point c, is common to
both series. This means that the simultaneous sounding
of a tone and its fifth will produce a disturbance at a
point corresponding to an octave below the lower tone
and corresponding also to a twelfth below the upper tone.
Hach of these tones alone produces a small disturbance at
this point and we have already seen that whether that
disturbance is or is not sufficient to produce stimulation
at this point depends upon the intensity of the sound. It
will also depend upon the intensity in this case, but if the
intensity of each be alone insufficient to produce this
effect when sounded alone, the two combined may still be
sufficient to produce this effect—an effect well-known to
musicians.

Similarly, any other two tones of sufficient intensity
may by their combined effects produce stimulation in the
organ of Corti at a point corresponding to a tone below
these two which is due to vibrations whose number is the
the greatest common measure of the vibration-numbers of
the two tones sounded. ‘This is also true even when the
two tones are not in harmony with each other, that is
when this resultant tone is more than an interval of a
seventeenth lower than the lower of the two tones. (‘The
ratio of numbers of vibrations of two tones separated by
an interval of a seventeenth is 1:5, ¢.g., A flat:e’.) And
this leads to the next problem.

Problem V. Jntra-cochlear effects of discords, consisting
of two tones.

348 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

It is hardly necessary to define a discord here; I will
use the term for any combination of two tones whose
beat-number is less than one fifth the vibration-number
of the lower tone, that 1s tones whose resultant tone is
removed from both of them by more than a major
seventeenth (2.e., more than two octaves and a major ©
third). }

If the interval be a major second (ratio of vibration-
numbers 8:9), the interval between c' and d' for instance,
then the highest resultant tone of the two is three octaves
below the lower of the two: 7.e., it 1s C,.

The basilar waves produced by combination of these
two are of complex form, eight waves of one series cor-
responding to nine of the other. At the points c’ and
d' and also at points between them the tectoria will be
brought down upon the sense-hairs, and with rhythmically-
varying force. The violence of the disturbance being at
its maximum at intervals of time corresponding to eight
periods of the lower tone (or nine of the upper): the
disturbance being at these moments about twice as
violent as that produced by either tone separately, while
at the moment intermediate between two of these the
disturbance will be practically nz.

At the point C,, every eighth wave of the series ¢’ and
every ninth wave of the series d’ will pass simultaneously
and there produce a stimulation leading to a sensation of
the tone C,.

An interesting example of this effect is Koenig’s, now
classic, experiment of sounding the notes c’” and d’” simul-

taneously (7.e., 2048 and 2304) the resultant (‘‘ differential’’)
~ tone 256 (c’) being distinctly heard. (See Nature, XLIL.,
p.- 190.)

Besides these resultant-tone stimuli there will be pro-
duced disturbances of the relative positions of tectoria

A NEW THEORY OF HEARING. 849

and sense-hairs at each point in the cochlea where a wave
of either series on the basilar passes a wave of the same or the
other series on the Reissnerian. Still considering the same
two tones c’ and d’ (which for convenience we will suppose
tuned to 256 and 288 vibrations per second respectively)
the secondary points of passings will correspond to the
following tones :—
Wepeeerics. f flat*, g', b' flat, d’,.g", a”, da’.
Padeerics. d;c, G, FP, D; OC, B, flat, A, flat, G,, F,,
cf etlab Ay fat* oD, C;.

feeeetics, {9¢ (—0 flat), 2c ~.¢ (=g9), 5c; xc’,
gsc’ (=e flat), ~.c’ (=d), r7e’, ec’ (=o),
etc.

In each series except perhaps the first, the intensity of
the disturbance would be greater at higher than at lower
points. In proportion to their height the effect would be
ereatest at the points in the second series. At points in
the first series the disturbance would occur twice in each
beat-period (,4, second) and in the following order :—1, 7,
eee, o, 4, 4, 5, 3, 6, 2, 7, 1.

The rhythm is more complex at the second series of
points and in the upper part of the series there is a
rhythmic variation of intensity at each. At the points in
the third series the intensity of disturbance is not so great
as in the other two and the rhythm is very complex.

Atc' and d@' there is a rhythmic variation in the intensity
of disturbance which in itself would produce an effect like
beats, 32 to the second.

How many of these points in the organ of Corti will
actually be stimulated will depend largely upon the
intensity of the primary tones employed.

It must be remembered that the signs now given, which
ordinarily stand for musical tones, here stand for the

(* An asterisk indicates that the point is indicated only approximately.)

" Pa
Ay ee

350 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

points in the organ of Corti corresponding to those tones,
and not for the tones themselves: the question of inter-
ference between them does not therefore arise. |

To which of these results the disagreeable and “restless”
effect of a discord is due cannot be said with certainty,
but it is probable the following all contribute to the
restlessness :

(1) The rhythmic variation of intensity of primary
stimuli beyond ordinary limits :

(2) The rapid succession of faint but unequal stimuli in
very widely separated regions of the organ, each stimulus
being due, in most cases, to two waves only and, hence,
giving rise to a comparatively ill-defined sensation :

(8) The probable stimulation of the whole area between
ce’ and d’.

With intervals less simple than this 8:9 discord the
beat-period becomes longer and the beats more distinct.
The ‘‘ beat”’ itself is the rhythmically recurring augmen-
tation of the primary stimuli: and their disagreeable effect _
is avoided in rapid music where the successive chords are
not sustained long enough to produce this effect.

It would seem to be inadvisable now to enter into
questions of more complex combinations of tones either
harmonious or discordant—apart even from limits of space.
The examples already given are sufficient to exemplify
the principles I have laid down.

An objection has been raised, first by myself and then
by almost everybody who has been led to discuss the
theory with me.

According to the theory now set forth, the region of the
cochlea where the stimulation by high tones occurs is

‘ = crim:
SS SRR

SS
‘

ee ae

== ‘a alieews
ha hate “eg oo nee ane
SS

near the apex; low ones producing a stimulation near
the base.
Older theories based on assumptions of ‘“‘ resonance”’ of

Se
= ‘ ‘

a

a REG NTO

A NEW THEORY OF HEARING. 351

of one part or another all locate the seat of stimulation by
high tones in the basal region and by low tones in the
apical. Pathological evidence, with the details of which
IT am not acquainted, shows that imjury—or at any rate
some injuries—to the apex of the cochlea lead to a
deafness to Jow tones, while the ear may still remain
sensitive to high ones.

The objection seems at first sight to be fatal but, if I
have rightly understood what is the nature of the patho-
logical evidence, it is not so.

A lesion of such kind as to destroy the elasticity of the
basilar membrane and thus prevent the passage of the
basilar wave to the apex of the spiral,‘ would itself
provide a new passage for the wave direct from basilar to
Reissnerian at the injured spot, and the whole course of
the cochlear wave would thus be shortened, the injured
' portion of the basilar serving as the turning-point of the
wave, serving in fact as a secondary helicotrema. Under
these circumstances waves of short period would still
“pass”? within the cochlea though not at the normal
point, and the whole course being shortened, waves of
long period would now fail to pass in the cochlea and
deafness to low tones would be the consequence.

In the earlier part of what has gone before, the attempt
to discover, by argument from physical considerations,
what changes of velocity of transmission the cochlear
wave undergoes in various parts of its length was abandoned
because the difficulties appeared insuperable. Having
now, however, seen the result of the physical consideration,
and being, as it seems, justified in concluding that the
stimulation of the nerve-ends is actually brought about by
the passing of waves on the two membranes, and the
resultant thrust of the tectorial membrane down upon the
sense-hairs we may use this physiological result as a means

352 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIHTY.

of answering the question which I was unable to answer
on purely physical grounds. - |

If the velocity remained constant in all parts it would
follow that the length of the region of the organ of Corti
set apart for stimulation by the various tones in the lowest

octave of audible sounds would be just half of the whole —

length of the organ. Above this region half the remainder
would serve for the next octave; and so on, each octave
having thus only one half as long an area as the preceding
one. The number of separate nerve-ends in each unit-
length of the organ is, however, very much the same in
all parts of the organ and the possibility of distinguishing
a difference of pitch between two tones depending pre-
sumably on their power of stimulating two different
nerve-ends, it follows that the accuracy with which we
can distinguish between two tones separated by only a

small musical interval would be enormously greater in the —

case of low tones than in the case of high ones. And this

is not the case. Very minute intervals, such as two or

three ‘cents,’ can be distinguished more easily in some
parts of the tone-scale than in others, but this difference
is exceedingly shght. It follows therefore that approxi-
mately equal lengths of the organ of Corti correspond to
approximately equal differences of pitch—equal that is in
the musical sense: or in other words the waves of the
basilar membrane must undergo an enormous acceleration
in their course along the membrane, the distance (in
space, not time) between two successive waves being thus
multiplied about 100-fold.

Near the upper limit of audibility there is a very marked
falling off in the power of discriminating between tones
differmg but little in pitch, and this means that this

acceleration of the wave is checked before the wave has

reached the very end of the organ of Corti.

=. °

*}

_——sA: NEW THEORY OF HEARING. 303

ll be noted that the physical limit of this acceler-
is not closely approached. That limit would be
approximately the velocity of sound in the liquid, %.e.,

nearly a mile per second, 1435 metres.

3
7

EXPLANATION OF PLATE XX.

Figs. 1 to 5. Diagrams, see pp. 328, 329, 337.

Fig. 6. Transverse section across middle canal of cochlea,
a 4 see p. 309. | .

| ‘Fig. 6a. Part of Fig. 6 enlarged.

REPORT on NEMERTINES observed at PORT
ERIN in 1894 and 1895.

By WoL, BEAvuMont, B.A.

EMMANUEL COLLEGE, CAMBRIDGE.

THE observations on which the present Report is based
were for the most part made during a stay of six weeks at
Port Erin in June and the earlier half of July, 1895. A
certain amount of attention had been previously devoted
to the group during portions of March and April, 1894.
My thanks are especially due to Prof. Herdman for
placing at my disposal the resources of the Biological
Station and for his ever ready help and encouragement,
and to Mr. T. H. Riches of Caius College, Cambridge, for —
very useful assistance and advice.

In Mr. Vanstone’s List (6) published early in last year
thirteen species of Nemertines were definitely reported as
occurring in the neighbourhood of Port Erin. With the
possible exception of the form therein recorded as Cari-
nella annulata, all of these have been observed by myself,
while eight additional species appear in the present
Report, viz.: Amphporus pulcher, A. dissemulans, Tetras-
iemma flavidum, Prosorhochmus claparedii, Micrura pur-
purea, M. fasciolata, M. candida, Cerebratulus fuscus.
The known Nemertine fauna of the district therefore —
numbers twenty-one, perhaps twenty-two, species; the
uncertainty depending on the identity of the species

appearing in the earlier Report as Carimella annulata, esi

Montagu. On the assumption which I believe to be

REPORT ON L.M.B.C. NEMERTINES. 355

i At ee re
: a =

correct, that the name is there used with the connotation
it bears in M‘Intosh’s Monograph, it is possible that the
-Nemertine so referred to is identical with that recorded
in the present Report as Carinella aragot, Joubin.

None of the parasitic species has yet been obtained in
the district ; no particular search was made for them by
myself beyond examining a few specimens of Galathea.
Malacobdella will probably be found; I had not, however,
an opportunity of obtaining that curious Nemertine, as
no living specimens of Cyprina islandica were dredged
during my visit.

Two cases of abnormality in the number of marginal
stylet sacs in the proboscis may be conveniently mentioned
here. Aspecimen of Tetrastemma candidum (of the deeper
water variety with white patch on head and dorsal white
line) was observed having three marginal sacs, while a
brownish yellow example of Tetrastemma dorsale had four
marginal stylet sacs, two on each side.

Before proceeding to deal with certain of the species in
detail it may be useful to give a complete list of the
Nemertines hitherto observed in the district.

Carinella aragot, Joubin. T. melanocephalum (Johnston).
? C. annulata (Montagu). T’. robertuane, M‘Intosh.
Cephalothria bioculata, Oersted.| Prosorhochmus claparedit, Ket.
: Amphiporus lactifloreus (Joh.). | Nemertes neesw, Oersted.

om

A. dissimulans, Riches. Lineus obscurus (Desor).

A. pulcher (Johnston). Lineus longissimus, Sowerby.
r Tetrastemma flavidum, Khrenb.| Micrura purpurea, J. Muller.
_ . dorsale (Abildgaard). M. fasciolata (Khrenberg).
| T. mgrum, Riches. M. candida, Burger.
— TL. immutabile, Riches. Cerebratulus fuscus (M‘Int.).
T. candidum (O. F. Muller). ? C. angulatus (O. F. Muller).
‘ T. vermiculatum (Quaitrt.).

- 856 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.
Order I—PROTONEMERTINI, Burger.
Family CARINELLIDA. |

Carinella aragot, Joubin.

C. annulata (pars), M‘Intosh, Hubrecht.
C. M‘intoshu, Burger, Riches.

This species has been dredged not infrequently this
year in about 15 to 18 fathoms on the shelly ground to
the north of the Halfway Rock, though it does not appear
to be abundant. The ground colour in the half dozen >
examples taken during my visit varied from dark chesnut
to chocolate. They were quite small, measuring only
from 2 to 7 cm. in length, but an exceptionally large
specimen over 20 cm. long in the preserved state was
shown to me by Mr. J. C. Sumner who had obtained it in
the same locality previous to my arrival. The Port Erin
form is exactly similar in external characters to examples
of this species seen by myself on the S.W. coast of
Ireland and at Plymouth. It 1s readily distinguished from —
Carinella annulata, Joubin, Burger, by the very charac-
teristic pale curved ciliated grooves on the dorsal surface
of the ‘‘ neck,” by the absence of a longitudinal white line
on the ventral surface of the body, and by the shape of
_ the head which is far less wide than in C. annulata; there
is considerably more white on the snout too in C. aragor
and the proboscis pore 1s more anterior in position.

2C. annulata (Montagu).

Mention has already been made of the possibility of the
species recorded as C. annulata in Mr. Vanstone’s Report
being identical with the last species. No example of the
Carinella annulata of Joubin and Burger has been met
with by myself at Port Erin; there is, however, no reason
why it should not occur in the district. ie

REPORT ON L.M.B.C. NEMERTINES. oot

Order I1.—_MESONEMERTINI, Burger.
Family CEPHALOTHRICIDA.

Cephalothriz bioculata, Oersted.

Besides being commonly found on the shore under
stones and among weeds, this species was occasionally
dredged near Port Erin in about 15 fathoms.

Order III.—_-METANEMERTINI, Burger.
Family AMPHIPORID.

Amphiporus pulcher (Johnston), M‘Intosh.

This well marked species was found to be fairly abundant
on the shelly ground off Port Erin in 15 to 20 fathoms;
it has also been dredged in the neighbourhood of Spanish
Head on several occasions.

The average length was 3 cm., the largest specimen
measuring 5 cm. All were of a uniform deep salmon
colour; no example with conspicuous gonads like the one
figured by M‘Intosh being met with. They were fre-
quently observed with the head telescoped into the anterior
part of the body as described by M‘Intosh, and the larger
ones swam vigorously when irritated. Amphiporus
pulcher is readily distinguished from A. dissitmulans and
A. lactifloreus not only by the possession of a reserve
central stylet and by the different position of the side-
organs but also by external characters: the broad oar-like
shape of the posterior half of the body, the large distinct
eyes not arranged in definite groups hke those of A.
lactifloreus, and especially the secondary grooves running
forward from the anterior cephalic furrows ; the. course of
the latter on the ventral surface is more transverse than
in the other two species referred to, so that they meet in
the middle line some distance behind the mouth-opening ;

- 858 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the secondary grooves are deep and well marked, far more
so than is indicated in M‘Intosh’s figure (Pl. XTV., fig. 11),
they are not confined to the ventral surface but extend
some distance on to the lateral portions of the dorsal
surface as well. 7 |

The figure and description of the cephalic furrows given
by Joubin (5, p. 129) may refer to Amphiporus dissimu- —
lans ; they certainly do not represent A. pulcher.

Amphiporus lactifloreus (Johnston).

Shore, common (see Vanstone).

Amphiporus dissimulans, Riches.

Numerous specimens of an Amphiporus closely re-
sembling A. lactifloreus, yet readily distinguishable from

that species on careful examination, were dredged in the

same localities as Amphiporus pulcher. No mature example
was taken ; the length of the majority was only from 6 to 9
mm. while a few reached 2to3cm. After the examination
of adult specimens of A. dissimulans at Plymouth I have
no hesitation in referring the Port Hrin form to this

species.

The larger specimens were brownish-yellow to dull
orange in colour, depending largely on the colour of the
alimentary tract and also on the presence of minute
sranules of yellowish brown pigment in the integument.
The smaller ones were of a fairly deep salmon colour due
entirely to the gut, no superficial pigment being as yet
apparent. The brain was plainly visible as a translucent
body of a pale yellowish tint.

The body was round and somewhat slender, and attained
its maximum thickness a little in front of the middle
point, diminution of size being very gradual in either
direction. 3

In the structure of the stylet apparatus and in the
position of the side-organs which open some distance in sj

REPORT ON L.M.B.C. NEMERTINES. 359

front of the brain this Nemertine agreed with Amphiporus
lactifloreus, but on a careful comparison of the two, certain
definite points of difference were invariably found. There
was a marked difference in the shape of the head, which
in A. lactifloreus is a rather long oval with the greatest
width just behind the lateral notches where the anterior
furrow passes from the dorsal to the ventral surface; in
A. dissumulans the head is more lozenge shaped, there
being a distinct angle at the widest part which is just an
front of the lateral notches of the anterior grooves; thence
the width decreases very distinctly as far as the lateral
notches formed by the posterior furrows, so that there is
a very distinct “‘neck.’”’ Another point of difference
concerns the arrangement of the eyes, which is very
characteristic in A. lactifloreus, where they are invariably
broken up into two distinct groups on each side, the
posterior group usually situated over the brain and
separated by a considerable interval from the anterior
series; while in A. dissimulans the eyes are arranged in
a continuous series on each side of the head, a few usually
extending inward towards the middle line in the region of
the anterior cephalic furrow. The smaller specimens at
Port Erin measuring 6 to 9 mm. in length usually had 8
to 10 eyes on each side; one 6 mm. long was noted having
as many as 14 eyes on each side, but this number was
generally found only in specimens of 2 to 3 cm. The
most posterior eye of the series was generally situated a
little in front of the brain, in no case were eyes observed
actually over the brain.

The figure given by Joubin to represent the head of
Amphliporus pulcher (5, p. 129) gives a fairly good idea of
the arrangement of the cephalic furrows in A. dissimulans ;
the anterior furrows usually bend more forward on the
ventral surface so as to run closer to and more nearly

360 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

parallel with the margin of the head than is there indicated,
and I have never been able to make out a branch groove
uniting the anterior and posterior furrows; at the angle
where the anterior groove bends forward, there is certainly
a branch given off, as in A. pulcher and other Enoplous
Nemertines, but it is quite short and ends abruptly.

In A. lactifloreus the arrangement is very similar; the
grooves being perhaps less conspicuous, and the anterior
grooves bending somewhat less forward on the ventral
surface so that the mouth-opening, which is in the usual
position close behind the tip of the snout, is a little in
front of the furrows and not intersected by them. Joubin’s
figure on p. 132 (5) does not appear to me to be at all an
accurate representation of the head of this species.

Whether the above characters are sufficient to entitle
Amphiporus dissumulans* to specific rank, or whether it
should be regarded merely as a variety of A. lactifloreus,
it is at any rate a well marked variety which appears to be
as strictly confined to the infra-laminarian zone as A.
lactifloreus to the littoral.

It remains to be mentioned that the small Port Erin
specimens differ somewhat from the adults seen at
Plymouth in respect to two features. In the former the
brain is not pink (this may be a juvenile character) and
secondly the head is less pointed and more distinctly
notched in front than is the case in the Plymouth form.

Family TETRASTEMMID&,

Tetrastemma flavidum, Ehrenberg.
This species appears to be well distributed in the

*It is almost needless to point out that Joubin is in error in assimilating
this species with A. pulcher. The two are very distinct, yet the possibility
of his having confused them is strongly suggested by the figure and accom-
panying description (5, p. 129) referred to above,

REPORT ON L.M.B.C. NEMERTINES. 361

neighbourhood of Port Erin but to be nowhere very
abundant. It has been obtained from the shore at Port
Erin and Perwick where it inhabits the smaller weeds on
reefs and in rockpools, and has also been dredged on
shelly ground in about 15 fathoms. All the specimens
taken were small, varying in length from 6 to 12 mm.
No mature individuals were met with, one immature
female being the only specimen in which genital organs
were observed.

In addition to the other well marked characters of this
species, the relative shortness of the proboscis may be
noted. ‘This organ extends back barely half the length of
the body from the anterior end.

The colour of the Port Hrin examples varied from
peach-colour to fairly bright orange red, and being due
almost entirely to the alimentary tract was much more
marked in the posterior half of the body. The fluid in the
proboscis-sheath had in some cases a pale yellow tint, as
had also the blood vessels in some specimens, more
especially in the posterior region of the body.

Tetrastemma dorsale (Abildgaard).

T'wo well marked colour varieties were found, apparently
restricted to different habitats.

The common dark brown type marbled with irregular
patches of reddish brown—narrow but of dark colour
where they cross the pale median dorsal band, widening
out but becoming paler on the sides of the body—is com-
monly found on the shore and in the Laminarian zone,
but does not appear to extend into the deeper water
outside. At all events on the shelly ground in 15—20
fathoms a much paler form of a brownish yellow colour
was exclusively found. In the absence of much superficial
pigment, the colour in these was due largely to the
alimentary tract. Some examples had hardly a trace of

362 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the reddish brown pigment so abundant in the shore form,
while others had obscure patches similar in disposition to
those present in the latter but much paler in colour.
Minute flakes of opaque yellowish white were scattered
all over the surface of the body as in the shore form, and
in many cases a concentration of these along the mid-
dorsal line produced a pale longitudinal streak. The
white flakes in this species are more superficial than the
reddish brown pigment. The latter is in the form of
much branched pigment cells whose processes run for the
the most part in the direction of the long axis of the body.

Tetrastemma nigrum, Riches. | |

A few examples of this species were met with both in
1894 and 1895. They were found in weeds from Port
Erin shore and from the Clets in the Calf Sound, and
were from 5 to 15 mm. in length.

Joubin, who does not appear to have met with this.
form, expresses doubt as to its being specifically distinct
from Tetrastemma dorsale. In shape, in the character of
its stylet apparatus and other anatomical details, it does
certainly agree with the latter, but so long as colour is
relied cn for the discrimination of the species of Tetras-
temma, it must be regarded as a well marked species,
much more so for example than are 7’. candidwm and its
allies.

In Tetrastemma mgrum the warm buff ground colour
is in most cases almost entirely concealed by black, or
almost black, reticular pigment, which is totally different
in character from anything seen in T’. dorsale; while on
the other hand the branched reddish brown pigment and
opaque white flakes so characteristic of the latter are
entirely absent in T’. mgrum.

Though Tetrastemma nigrum varies considerably, the
variation met with merely concerns the extent to which

REPORT ON L.M.B.C. NEMERTINES. 363

the surface of the body is covered with black pigment,
and in no way tends to bridge the gap between the type
and 7’. dorsale. In a few individuals the ground colour
is nowhere visible except along a narrow irregular strip of
the ventral surface, usually, however, there is a longitudinal
median dorsal band free from black pigment, varying
much in width in different individuals and frequently
interrupted by one or more bridges of pigment; while one
or two examples have been met with in which the black
pigment was confined to a series of isolated more or less
rounded patches, the aggregate area of which was less
than that of the intervening pale surface.

Tetrastemma immutabile, Riches.

Two Nemertines agreeing in shape and colour with the
description given by Riches were found in 1894 in material
from the Clets, and a couple more turned up in shore
material from Perwick in 1895. They were from 8 to 6
mm. long.

No intermediate varieties connecting this form with
Tetrastemma dorsale were met with. The median dorsal
band consists of branched reddish brown pigment cells
the processes of which tend to run longitudinally, and
also of round granules of darker brown colour, a few of
which are scattered over the rest of the dorsal surface.
In none were genital organs observed.

Tetrastemma candidum (O. F. Muller).

This species was found in considerable abundance
inhabiting the weeds between tide-marks at Port Erin,
Perwick and Port St. Mary. Very few specimens exceeded
1 cm. in length; the majority were considerably less, yet
many even of the smaller ones had well developed genital
organs, one male not more than 4 mm. long having 6 pairs
of testes containing ripe spermatozoa.

These littoral forms, though varying a good deal in colour

- 864 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

and other details, agreed for the most part in having the
head more rounded in outline than is the case in Tet-
rastemma melanocephalum and in this respect differed
slightly from the deeper water form described below. In
colour the majority were of various shades of yellow—
from pale straw to deep apricot-yellow—flesh-coloured,
yellowish brown and dull orange varieties were also
observed while a few were of a greenish-yellow, the pre-
vailing yellow tint being modified by the green colour of
the alimentary tract, but no specimen was met with at
all resembling the bright green type figured by M‘Intosh.

No example of this shore form exhibited any trace of
opaque white either on the head or in the form of a
median dorsal line. Only two individuals were found
showing a tendency by the development of brown pigment
on the head to connect this species with the two following
ones. Both were mature females agreeing in other
respects with the more typical specimens of 7’. candidum
from the same source. In one, about 1 cm. long, there
was a band of brown pigment between the anterior and
posterior eyes on each side but not quite reaching the
latter; in the other an obscure brown patch extended
across the head immediately behind the anterior pair of
eyes, the colour being more pronounced between the
anterior and posterior eyes on each side.

In the deeper water (15 to 20 fathoms) outside Port Erin
on shell and coralline bottom, a rather well marked form
was found, which may be described here since it lacks
the characteristic markings of the two following species.
It is, however, quite a debateable point whether it ought
not rather to be added to the series formed by Hubrecht’s
species 7’. melanocephalum, T. coronatum and T. diadema
as an extreme variety of T. melanocephalum. It seems
to come very close to 7’. diadema but in none of the many

REPORT ON L.M.B.C. NEMERTINES. 365

specimens examined was there any tendency to the devel-
opment of brown pigment on the head.

Most of the specimens taken measured from 1 to 2 cm.
in length; many had well developed genital organs. In
shape they resembled TJ’. melanocephalum, the more
spathulate head distinguishing them from the littoral form
dealt with above. The colouring was fairly constant ;
the majority were a more or less deep apricot-yellow, a
few were of a rather more greenish or brownish tint and
one flesh-coloured individual was noted. In all a con-
spicuous oblong patch of opaque white flakes was placed
transversely between the anterior and posterior pairs of
eyes, from which in most cases a median longitudinal line
of similar white flakes extended back along the dorsal
surface, generally the whole length of the body. The
eyes were black, well defined usually and about equal in
size, the posterior pair being often somewhat closer
together than the anterior in the well extended animal.

In the character of the proboscis they agreed with the
more typical T. candidwm from the shore rather than
with T. melanocephalum, the stylet of the central appara-
tus being on the whole longer in proportion to the handle
than is the case in the latter. Much stress cannot,
however, be laid on this point, for the stylet apparatus,
especially the handle, varies so much in form as to be of
little value in discriminating between such closely allied
species as 7’. candidum, T. vermiculatum and T’. melano-
cephalum, though sufficing as a general rule to distinguish
these three as a group from other Enoplous Nemertines.

Mention may here be made of two or three brownish
yellow Nemertines agreeing fairly well in general appear-
ance with T'etrastenuna candidum, but having the handle
of the central stylet very long, its posterior border being
straight with rounded corners, thus resembling in outline

866 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

the central apparatus of some of the Port Evin specimens
of Prosorhochmus. The examination of more specimens
is desirable before forming any conclusion as to their
relation with other forms.

Tetrastemma vernaculatum (Quatrf.), M‘Intosh.

This usually abundant species does not seem to be at
all common in the district. I have only met with three
specimens. ‘T’wo of a deep flesh-colour—the larger a
mature female 12 mm. long—were found in weeds from
between tide-marks at Perwick, the third came from the
Clets in Calf Sound and was of a rich apricot-yellow
colour. In all three the head was fairly oval in outline,
a point which serves in some degree to distinguish this
from closely allied species, but in none was there a trace
of the dorsal white line which Joubin considers its most
reliable character.

Tetrastemma melanocephalwm (Johnst.), M‘Intosh.

T. melanocephalum, Hubrecht.
T. coronatum, Hubrecht.

This species is fairly abundant and well distributed in
the district, inhabiting the weeds between tide-marks and
also the shell and coralline ground in 15 to 20 fathoms.
At Easter, 1894, a few large greenish specimens were
found (up to 33} cm. long), but the numerous examples
collected in June and July, 1895, were all immature,
averaging only about 1 cm. in length. ‘They presented a
considerable range of colour-variation, different shades of
flesh and salmon-colour and of yellow from pale greenish-
yellow to deep apricot being noted as the ground colour.

A median dorsal white line was present in many
specimens, and the majority had more or less opaque
white on the head in front of, and immediately posterior
to, the patch of dark pigment. The latter was reddish
brown or chocolate in colour and very variable in shape.

REPORT ON L.M.B.C. NEMERTINES. 367

In many cases it was a deep crescent, the horns barely
touching the anterior pair of eyes and the convex posterior
margin extending to about halfway between them and
the posterior pair. ‘This variety appears to correspond to
Hubrecht’s species 7’. coronatwm. In others again an
oblong band of pigment extended quite across the head
(almost reaching its lateral margins) immediately behind
the anterior pair of eyes and covering one-half to two-
thirds of the space between them and the posterior pair.
This seems to be Hubrecht’s typical T. melanocephalum.
With respect to these two forms, however, my expericnce
both at Port Erin and elsewhere coincides with that of
Riches, a complete series of intermediate varieties making
it impossible to regard them as more than slightly divergent
members of one variable species.

This opinion was at one time held by Joubin too, but
in his later work (5) 7’. coronatum is separated from T.
melanocephalum and united with T. diadema, Hubrecht.
No Nemertine exactly agreeing with the latter has been
met with by myself, but the Nemertine from the coralline
zone described under the head of 7. candiduwm seems to
come very near it. That form, however, did not show
any tendency to vary in the direction of 7’. melanocephalum
so far as my observations went.

A character given for the present species by M‘Intosh:
“Marginal stylets-sacs placed considerably in front of the
central apparatus’’—has been repeated by other authors,
though to what extent this point has been confirmed by
their own observations does not appear. Ihave examined
the stylet apparatus in many individuals and have so far
failed to find a single case exhibiting this character, or
any difference in this respect between 7’. melanocephalum
and its allies.

A Nemertine having the form and general appearance

368 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of T. melanocephalum was observed, which resembled a
specimen described by Riches in having the dark pigment
patch broken into two by a space in the middle line free
from pigment, and thus connecting the present spear
with 7. vermiculatum.

Tetrastemma robertiane, M‘Intosh.

Two specimens of this rare species were dredged in
1895 on the shelly ground off Port Erin in about 15
fathoms. Lochmaddy and Bressay Sound where it was —
dredged by M‘Intosh appear to be the only other localities —
where it has been found.

The two examples observed by myself agreed fairly well
with M‘Intosh’s figures and description, though in neither
was the head so pointed as he represents it 1n the woodcut
on page 167 of the Monograph. The central stylet
apparatus did not differ materially from his figure (p. 65).

The larger individual, about 1 cm. long, had the median
dorsal white line, the reddish brown lateral stripes and
collar, the latter not quite encircling the neck ventrally. _
There was a patch of opaque white flakes between and in
front of the posterior eyes which were much smaller than
the anterior pair. The smaller one was similar except
that the brown collar was wanting.

A Nemertine from the same locality ressHibnee the
above in shape but differing somewhat in arrangement of
colour and also in the character of its stylet apparatus
may be conveniently described here. Like them it was
exceedingly active. It measured about 8 mm. in length.
The ground colour was apricot-yellow, a large patch of
opaque white flakes extended across the dorsal surface of |
the head between the anterior and posterior eyes, a little
behind the latter was a chocolate brown collar not quite
complete ventrally, and from this a median longitudinal
band of the same colour extended back the whole length

REPORT ON L.M.B.C. NEMERTINES. 369

_ of the dorsal surface. The anterior eyes were crescentic
in shape and twice the size of the more rounded posterior
pair.

The central stylet apparatus was very large, the stylet
somewhat longer than was the case in the two typical
specimens of T’. robertiane, and the handle very long, its
posterior border straight with rounded corners.

Prosorhochmus claparedw, Keferstein.

The Nemertine referred here was met with not un-
commonly among material dredged on the shell ground
lying to the north of the Halfway Rock in 15 to 20 fathoms.
There can be little doubt that it is identical with the
Plymouth form described by Riches under this name,
though I failed to make out a superior lobe on the head.
Both differ very considerably in general appearance from
the figures of Prosorhochmus claparedw given by M‘Intosh,
and ought very possibly to be regarded as a distinct species.

In the Port Erin specimens, which were from 8 to 12
mm. long, the body was stout and rounded, tapering from
the middle towards either end. The head broad, rounded
in outline and with pronounced notch in front, very
convex dorsally, and well marked off by. conspicuous
cephalic furrows resembling those of Tetrastemma can-
didwm in disposition. The colour was a dull brownish
orange resulting from the deep apricot yellow colour of
the alimentary tract modified by a sprinkling of minute
granules of superficial reddish brown pigment all over
the body but much more plentiful on the dorsal surface.
In one or two specimens there was a slightly greater
development of pigment between the anterior and posterior
cephalic grooves, but to nothing lke the extent seen in
the Plymouth form. The greater portion of the dorsal
surface of the head in front of the anterior furrows was
covered with opaque white flakes; similar flakes were

370 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

usually disposed as a fan on the posterior extremity of

the body, while in one individual minute white grains
superficial to the brown pigment appeared all over the
body. Inno case was there any tendency to the formation
of a dorsal white line.

The eyes were reddish brown and crescentic in shape,

the ragged concave side being directed outward. The
anterior pair were much larger than the posterior, and
the latter were much closer together than the former
when the animal was well extended; the two pairs, how-
ever, were not approximated as represented in tue figures
of M‘Intosh but on the contrary were separated by an

interval quite as great as in the case of T'etrastemma can-

didum for instance.

The central stylet apparatus agreed precisely with the
figure given by M‘Intosh (Pl. XIII, fig. 1), except in two
specimens wherein the handle of the central stylet was
long and with a straight posterior border as in the others,
but the corners were rounded off instead of sharp.

A number of females containing numerous large ova
were seen but in no case was there any sign of commencing

development.
Family NEMERTIDA,

Nemertes neesw (Oersted).

A young individual 8 cm. long dredged in about 20
fathoms off Peel on June 28rd, 1895, is the only example
of this species which has so far as I know been taken
below low water mark in the district. Large specimens
were commonly found on the shore in all suitable localities.

Order IV.—HETERONEMERTINI, Burger.
Family LINEID2.

Lineus obscurus (Desor).
Shore, common (see Vanstone).

t q

REPORT ON L.M.B.C. NEMERTINES. ayia

Lineus longissimus, Sowerby.

This species was not met with at all commonly; a few
specimens were found at low tide at Port Erin in 1894,
and it was dredged off Bay Aldrick in about 17 fathoms
in 1895.

Micrura purpurea (Dalyell), J. Muller.

Two examples of this species were dredged in about 15
fathoms to the north of the Halfway Rock on shelly
bottom. They were 6 to 7 cm. long, and in colour rich
purple-brown with the characteristic patch of yellow on
the snout. No eyes could be made out in either.

Micrura fasciolata, Khrenhberg.

This species was dredged not infrequently in 15 to 20
fathoms on the shell ground near Port Erin, but can
hardly be considered abundant. Length 3to6 cm. In
addition to specimens of the typical brownish red colour
with conspicuous white bands, a few were met with
resembling a variety described by M‘Intosh; these were
ereenish and greyish brown in colour and the white bands
were very obscure, or even entirely absent, so that, but
for the possession of a caudal appendage, they bore a very
close resemblance to similarly coloured varieties of Lineus
obscurus.

An individual of this species, when accidentally dropped
into a solution of picric acid, disgorged a Polychaete
worm about half its own length.

Micrura candida, Burger.

Cerebratulus lacteus, Hubrecht.

An opaque white Nemertine about 4 cm. long dredged
in June, 1895, to the north of the Halfway Rock in about
15 fathoms, proved to belong to the above species. It was
readily distinguished from pale examples of Cerebratulus
fuscus by its rounder more attenuated body and less
pointed head, and by the absence of eyes. ‘The surface

372 TRANSACTIONS LIVERPOOL BIOLOGICAL SOCIETY.

of the body was much wrinkled transversely, the intervening ~
folds being densely crowded with flakes of opaque white,
so that the brownish flesh-colour of the deeper tissues was —
only visible in the wrinkles. No granular brown pigment
was present. The brain showed as an ill defined pinkish
mass. A caudal appendage was present. A second
specimen 6 mm. long was dredged on the same ground
later. The ground colour was white with a yellowish
tinge due to the gut, the wrinkling was less marked than
in the larger individual, which it otherwise closely
resembled. The single specimen taken by Riches at
Plymouth is the only previous British record for this
species.

Cerebratulus fuscus (M‘Intosh), Hubrecht.

The posterior end of a large individual of this species
was dredged in the neighbourhood of Spanish Head on
the ‘‘ Lady Loch” excursion of June Ist, 1895. The
dorsal surface was somewhat profusely spotted with brown
pigment; this and the very characteristic shape left no
doubt as to its identity. .
~ During the course of the same month one specimen
about 2 cm. long and some half dozen smaller ones were
obtained off Port Erin in 15 to 20 fathoms.

REFERENCES.

1. M‘IntosH.—A Monograph of British Annelids. Pt.
I., Nemertines. eats

2. HuBRECHT.—Genera and Species of European Nemer-
tines. Notes from Leyden Museum, XLIV.,1879.

3. BurcER.—Zur Systematik der Nemertinenfauna des ¥ 4
Golfes von Neapel. Nachrichten v. d. K. Gesell-
schaft d. Wissenschaften zu Gottingen, 1892.

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REPORT ON L.M.B.C. NEMERTINES. 373

4. RicHes.—The Nemertines of Plymouth Sound. Journal
of the Marine Biological Association, 1893.

5. JouBIn.—Faune France: Les Némertiens, 1894.

6. VANSTONE.—The Nemertines of Port Hrin, Isle of Man.
Trans. Liverpool Biological Society, 1894.

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