^Harvard University
Dbrary of
Che CDedical School
and
Ghe School of ‘Public ^Health
ERRATA
Vol. XIV, No. 3, p. 342, Table I: for 103 : 19 3 read 108 : 20 5 ;
and p. 343, Table III : for 269 : 39 3 read 265 : 387.
Annals of Tropical Medicine
and Parasitology
THE UNIVERSITY OF LIVERPOOL
Annals
OF
Tropical Medicine and
Parasitology
ISSUED BY THE
Liverpool School of Tropical Medicine
Edited by
Professor J. W. W. STEPHENS, M.D.Cantab., F.R.S.
Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S.
Professor WARRINGTON YORKE, M.D.
Professor B. BLACKLOCK, M.D.
VOLUME XV
(April 27,1931, to December 30,1901)
With Frontispiece, thirty-one plates, one hundred and twenty-seven
figures in text, five maps, and four charts
LIVERPOOL:
AT THE UNIVERSITY PRESS, 57 ASHTON STREET
CONTENTS
» . j. . r
No. i. April 27, 1921
Allmand, Dorothy. -
Liverpool School of Tropical Medicine. Scientific Record ... ... . ...
Breinl, A.
- On the ‘ Arneth Count * in Hookworm-Infected White Children in North
Queensland .
Macfie, J. W. S.; and Ingram, A.
Bronchomoniliasis Complicating Pulmonary Tuberculosis in a Native of the
Gold Coast, West Africa .
Blacklock, B.
Notes on a Case of Indigenous Infection with P. falciparum ....
Blacklock, B.; and Carter, H. F.
Observations on Mosquitoes in the Isle of Man ... ..1 .
Hill, G. F.
Notes on Some Unusual Breeding-places of Stegomyia fasciata , Fabr., in
Australia. ...
Hill, G. F.
. .. Musca domestic Linn., as a ‘ Bush Fly * in Australia .
Newstead, R.; and Evans, Alwen M. . ..
New Tsetse-Flies ( Glossina) from the Belgian Congo ... ..
Newstead, R.; and Sinton, J. A.
.On a Collection of Pappataci Flies ( Phlebotomus ) from India ... ...
PAGE
1
49
53
59
73
9i
93
95
103
v
CONTENTS
No. 2. July 16, 1921
SlNTON, J. A. PAGE
Notes on Oriental Sore in Russian Turkestan and the Results of Treatment
with Iiyections of Tartar Emetic Solution. 107
Barret, H. P.
A Method for the Cultivation of Blastocystis . 113
Scott, Henry Harold
The Incidence of Intestinal Parasites, especially with regard to the Protozoa,
amongst Symptomless Carriers in Jamaica . . 117
Scott, Henry Harold .
A Study of the Sizes of Entamoeba histolytica Cysts amongst Symptomless
Carriers in Jamaica . . 133
Scott, Henry Harold
A Study of the Sizes of Entamoeba coli Cysts amongst Symptomless Carriers in
Jamaica. 149
Southwell, T. *
Cestodes from Indian Poultry. 161
Southwell, T.
Cestodes from African Rats . 167
Southwell, T.
A New Species of Cestoda from a Cormorant . 169
Stephens, J. W. W.; and Adler, S.
A Case of Suspected Leprosy. 173
vi
CONTENTS
No. 3. September 30, 1921
Carter, Henry F.; Ingram, A.; and Macfie, J. W. S.
Observations on the Ceratopogonine Midges of the Gold Coast with Descriptions
of New Species. Part IV .
Scott, Henry Harold.
The Prevalence and Character of Tuberculosis in Hongkong. Part I
Scott, Henry Harold.
The Prevalence and Character of Tuberculosis in Hongkong. Part II
Kbnnan, R. H.
Multiple Aneurysms in a Child .
Yorke, Warrington ; and Southwell, T.
Lappeted Anoflocephala in Horses.
Gordon, R. Montgomery ; and Young, C. J.
The Feeding Habits of Sugomyia calopus , Meigen .
Yorke, Warrington ; and Adler, S.
Note on a Case of Leprosy.
Macfie, J. W. S.
Notes on some Fungal Infections in West Africa. .
Macfie, J. W. S.; and Ingram, A.
A Fungus of the genus Nocardia Cultivated from Heart Blood
Newstead, R.; and Evans, Alwen M.
Report on Rat-Flea Investigation..
Young, C. J.
Natural Enemies of Sttgemyia caUpvs, Meigen .
vii
page
177
213
227
*45
249
265
269
271
283
287
301
PAGE
CONTENTS
No. 4. December 30, 1921
Ingram, A.; and Macfie, J. W. S. ' ^
West African Ceratopogoninae . 313
Ma£fie, J. W. S.
The Effect of Saline Solutions and Sea-water on SUgomyia jasciata . 377
Scott, Henry Harold.
The Prevalence and Character of Tuberculosis in Hongkong. Part III ... 381
Ihle, J. E. W.
On the Genus Cylicostomum . 397
Maplestone, P. A.
Notes on Australian Cestodes. Part I . 403
Maplestone, P. A.
* Notes on Australian Cestodes. Part II ... .. ... ... 407
Maplestone, P. A.
Notes on Ulcerative Granuloma . 4*3
MacGregor, Malcolm E.
The Structural Differences in the Ova of AnopheUs maculipennis , A. bifurcatus
and A . plumbeus . . ... ... 417
Adler, S.
The Trypanocidal Effect of Phenylglycine Amido Arsenate of Sodium on
T. brucei in Rats and T. rhodesiense in Mice.; ... ... 427
Adler, S.
Note on Bismuth as a Trypanocide.. 433
Stephens, J. W. W.
Malaria on a Venezuelan Oilfield. . ... ... 435
Evans, Alwen M.
Notes on Culicidae Collected in Venezuela . 445
Southwell, T.; and Maplestone, P. A.
A Note on the Synonymy of the Genus Zschokkeella , Ransom, 1909, and of the
Species Z. guitieensis (Graham, 1908). . 455
Davey, J. B.; and Newstead, R.
Mosquitoes and other Blood-sucking Arthropods of the Upper Shiri River,
Nyasaland ... ... ‘ ... . .- ... .:. 457
Blacklock, B.
Breeding Places of Anopheline Mosquitoes in Freetown, Sierra Ledne... ...* 463
Blacklock, B.
Notes on an Apparatus for the Individual Breeding of Mosquitoes . 473
Yorke, Warrington. _ ■ .
The Treatment of a Case of Rhodesian Sleeping Sickness by the Preparation
known as ‘ Bayer 205 , . . . 479
viii
Volume XV
April, 1921
No. 1
ANNALS
or
TROPICAL MEDICINE AND
PARASITOLOGY
ISSUED BY
THE LIVERPOOL SCHOOL OF TROPICAL MEDICINE
Edited by
Professor J. W. W. STEPHENS, M.D., Cantab., F.R.S.
Professor R. NEWSTEAD, M.Sc., J.P., F.R.S., A.L.S., F.E.S., Hon. F.R.H.S.
Professor WARRINGTON YORKE, M.D.
Professor B. BLACKLOCK, M.D.
THE MARY KINGSLEY MEDAL
This medal was struck in commemoration of the work of the late
Miss Mary Kingsley in West Africa, and is conferred in recognition of
distinguished scientific achievement.
HONORARY RECIPIENTS
Her Royal Highness Princess Christian
Lord Lister
The Right Hon. Joseph Chamberlain
Prince Auguste d’Arenberg
Mrs. Pinnock
Mr. William Adamson
Professor William Carter
RECIPIENTS
/ 905 -
Colonel Sir David Bruce, K.C.B.
Geheimrath Professor Robert Koch
Dr. A. Laveran
Sir Patrick Manson, K.C.M.G.
1907 —
Professor Danielewsky
Dr. Charles Finlay
Mr. W. M. Haffkine
Professor Golgi
Colonel Gorgas
Professor Theobald Smith
1910 —
Sir William Macgregor, G.C.M.G.
Professor R. Blanchard
Dr. Anton Breinl
Professor Angelo Celli
Dr. C. W. Daniels
Surgeon-General Sir Alfred Keogh
Colonel W. G. King
Professor Nocht
Professor G. H. F. Nuttall
Major Leonard Rogers
Professor J. L. Todd
Surgeon-General Walter Wyman
1913 —
Professor Fred V. Theobald
1917 —
Dr. Griffith Evans
1919 —
Dr. J. W. Scott Macfie
The Oswaldo Cruz Institute, Rio de
Janeiro
1920 —
Major E. E. Austen, D.S.O.
Dr. A. G. Bagshawe, C.M.G.
Dr. Andrew Balfour, C.B.
Dr. A. L. G. Broden
Mrs. Chalmers, in recognition of the
work of the late Dr. A. J. Chalmers
Professor B. Grassi
Professor R. T. Leiper
Professor F. Mesnil
Dr. Edmond Sergent
Dr. C. W. Stiles
Dr. T. Zammit
THE INCORPORATED
LIVERPOOL SCHOOL OF TROPICAL MEDICINE
Founded by Sir ALFRED LEWIS JONES, K.C.M.G
{Affiliated with the University of Liverpool)
Hon. President : H.R.H. Princess Christian
(Princess of Great Britain and Ireland)
Chairman : Sir Francis C. Danson
Vice-Chairman : Professor R. CATON, C.B.E.
Hon. Vice-Presidents : The Earl of Derby, K.G.
Viscount Milner, G.C.B.
Lord Pirrie, K.P.
Lord Lbverhulmb
Sir Edward Merbwbther, K.C.V.O.
Sir Owen Philipps, K.C.M.G.
Mr. O. Harrison Williams
COMMITTEE
Sr H. J. Read, K.C.M.G. Colonial Office
Vice-Chancellor J. G. Adami, F.R.S. University of Liverpool
Professor R. CATON, C.B.E.
Mr. H. Wade Deacon
Professor J. M. Beattie
Professor Dakin
Mr. E. G. Buckley
Mr. T. Woodsend
Mr. T. F. Harrison
Mr. A. R. Marshall
Mr. W. Roberts
Mr. R. B. Miller
Mr. J. W. Alsop, O.B.E.
Mr. G. Brocklehurst
Mr. H. D. Dickie
Professor E. GLYNN
Professor W. Herdman, C.B.E., F.R.S.
Professor E. W. Hope, O.B.E.
Mr. David Jones
Mr. J. Pickering Jones
Mr. J. A. Tinne
Captain R. Rankin, Hon. Treasurer
Secretary,
H 24—25, Exchange Buildings, Liverpool.
| Council of University of Liverpool
Senate of University of Liverpool
Royal Southern Hospital
Steamship Owners' Association
■ Shipowners' Association
West African Trade Association
Staff, 1921
Professors - - JOHN WILLIAM WATSON STEPHENS, M.D.,
Can lab., F.R.S. Alfred Jones Professor of
Tropical Medicine
ROBERT NEWSTEAD, J.P., F.R.S., M.Sc.,
A.L.S., FrE.S., Dutton Memorial Professor of
Entomology
WARRINGTON YORKE, M.D., Walter Myers
Professor of Parasitology
B. BLACKLOCK, M.D., D.P.H., Professor of
Tropical Diseases of Africa
Lecturers - - ALWEN M. EVANS, M.Sc., Assistant Lecturer on
Entomology
Prof. E. W. HOPE, M.D., D.Sc., Lecturer on
Municipal Sanitation
R. H. KENNAN, M.D., Lecturer on Tropical
Sanitation
A. W. NOEL PILLARS, F.R.C.V.S., Honorary
Lecturer on Clinical Veterinary Parasitology
T. SOUTHWELL, A.R.C.Sc., F.Z.S., Assistant
Lecturer on Parasitology
Honorary Statistician WALTER STOTT
Honorary Lecturer in
Tropical Sanitation WILLIAM THOMAS PROUT, M.B., C.M.G.
Royal Infirmary, Liverpool
Physician - - JOHN WILLIAM WATSON STEPHENS, M.D.,
Cantab., F.R.S.
Assistant Physician - WARRINGTON YORKE, M.D.
House Physician and
Clinical Pathologist S. ADLER, M.B., Ch.B.
The Mangos Research Laboratory
Director - - HAROLD WOLFERSTAN THOMAS, M.D.,
C.M.
Research Assistants - RUPERT MONTGOMERY GORDON,
M.B., Ch.B.
CHARLES JAMES YOUNG, M.B., Ch.B.
NOTICE
The following courses of instruction will be given by the
Liverpool School of Tropical Medicine during 1921 :—
Full Course began 10 January. Advanced Course begins 1 June.
Diploma Examination, 14 March. Certificate Examination, 1 July.
Full Course begins 19 September.
Diploma Examination, 12 December.
These dates are subject to revision.
The full Course of Instruction is'open to all qualified medical men,
and the Examination to all students who have taken out this full course.
Fee for the full Course of Instruction—Thirteen Guineas.
Fee for the Diploma Examination—Five Guineas.
Fee for the Short Course of Instruction—Four Guineas.
Fee for the use of a School microscope during one term—One Guinea.
For prospectus and further information, application should be made
to the Dean of the Medical Faculty, University of Liverpool.
The following have obtained the Diploma in Tropical Medicine of
the University of Liverpool :—
Diploma in
Data if
Diploma
1904 Augustine, Henry Joshua
1904 Bennett, Arthur King
1904 Bruce, William James
1904 Byrne, John Scott
1904 Clayton, Thomas Morrison
1904 Dalziel, John McEwen
1904 Dee, Peter
1904 Greenidge, Oliver Campbell
1904 Hehir, Patrick
1904 Khan, Saiduzzafor
1904 Laurie, Robert
1904 Maclurkin, Alfred Robert
1904 McConnell, Robert Ernest
1904 Nicholson, James Edward
1904 Phi lip ton, Nicholas
1904 Sharman, Eric Harding
1904 Thomson, Frank Wyville
1904 Walker, George Francis Clegg
1905 Anderson, Catherine Elmslie
1905 Brown, Alexander
1905 Caldwell, Thomas Cathcart
1905 Critien, Attilio
1905 Hoot on, Alfred
1905 Hudson, Charles Tilson
1905 Illington, Edmund Moritz
Tropical Medicine
Daii if
Diploma
1905 Macfarlane, Robert Maxwell
1905 Mad dock, Edward Cecil Gordon
1905 Moore, James Jackson
1905 Nightingale, Samuel Shore
1905 Rad cliff e, Percy Alexander Hurst
1905 Young, John Cameron
1906 Adie, Joseph Rosamond
1906 Arnold, Frank Arthur
1906 Bate, John Brabant
1906 Bennetts, Harold Graves
1906 Carter, Robert Markham
1906 Chisholm, James Alexander
1906 Clements, Robert William
1906 Dundas, James
1906 Faichnie, Norman
1906 Jeffreys, Herbert Castelman
1906 Mackenzie, Donald Francis
1906 Pailthorpe, Mary Elizabeth
1906 Palmer, Harold Thornbury
1906 Pearse, Albert
1906 Sampey, Alexander William
1906 Smithson, Arthur Ernest
1906 Taylor, Joseph van Soraeron
1906 Taylor, William Irwin
1906 Tynan, Edward Joseph
Data of
Diploma
1906 Watson, Cecil Frandi
1906 Willcoclu, Roger Durant
1906 Williamson, George Alexander
1907 Allan, Alexander Smith
1907 Allwood, James Aid red
1907 Bond, Ashton
1907 Branch, Stanley
1907 Collinson, Walter Julius
1907 Davey, John Bernard
1907 Donaldson, Anson Scott
1907 Fell, Matthew Henry Gregson
1907 Gann, Thomas William Francis
1907 Graham, James Drummond
1907 Hiscock, Robert Carroll
1907 Keane, Joseph Gerald
1907 Kennan, Richard Henry
1907 Kenrick, William Hamilton
1907 Le Farm, George Ernest Hugh
1907 Mackey, Charles
1907 Maddox, Ralph Henry
1907 McCarthy, John McDonald
1907 Raikes, Cuthbert Taunton
1907 Ryan, Joseph Charles
1907 Vallance, Hugh
1908 Caverhill, Austin Mack
1908 Crawford, Gilbert Stewart
. 1908 Dalai, Kaikhusroo Rustomji
1908 Dansey-Browning, George
1908 Davidson, James
1908 Dickson, John Rhodes
1908 Dowd all, Arthur Melville
1908 Glover, Henry Joseph
1908 Greaves, Francis Wood
1908 Good body, Cecil Maurice
1908 Harrison, James Herbert Hugh
1908 Joshi, Lemuel Lucas
1908 Le Fanu, Cecil Vivian
1908 Luethgen, Carl Wilhelm Ludwig
1908 Mama, Jamshed Byramji
1908 McCay, Frederick William
1908 McLellan, Samuel Wilson
1908 Pearce, Charles Ross
1908 Schoorel, Alexander Frederik
1908 Smith, John Macgregor
1908 Stewart, George Edward
1908 Tate, Gerald William
1908 Whyte, Robert
1909 Abercrombie, Rudolph George
1909 Allin, John Richard Percy
1909 Armstrong, Edward Randolph
1909 Barrow, Harold Percy Waller
1909 Beatty, Guy
1909 Carr-White, Percy
1909 Chevallier, Claude Lionel
1909 Clark, William Scott
1909 Cope, Ricardo
1909 Fleming, William
1909 Hanschell, Hother McCormick
1909 Hayward, William Davey
1909 Henry, Sydney Alexander
1909 Innes, Francis Alexander
1909 Jackson, Arthur Frame
1909 Kaka, Sorabji Manekji
Dots §f
Diploma
1909 McCabe-Dallas, Alfred Alexander
Donald
1909 Meldrum, William Percy
1909 Murphy, John CulEnan
1909 Samuel, Mysore Gnananandaraju
1909 Shroff, Kawasjee Byramjee
1909 Thornely, Michael Harris
1909 Turkhud, Violet Ackroyd
1909 Webb, William Spinks
1909 Yen, Fu-Chun
1910 Brabazon, Edward
1910 CastelHno, Louis
1910 Caulcrick, James Akilade
1910 Dowden, Richard
1910 Haigh, William Edwin
1910 Hamilton, Henry Fleming
1910 Hefferman, William St. Michael
1910 Hipwell, Abraham
1910 Homer, Jonathan
1910 Houston, William Mitchell
1910 James, William Robert Wallace
1910 Johnstone, David Patrick
1910 Korke, Vishnu Tatyaji
1910 Macdonald, Angus Graham
1910 Macfie, John Wm. Scott
1910 Manuk, Mack Walter
1910 Murison, Cecil Charles
1910 Nanavati, Kish a vial Balabhai
1910 Nauss, Ralph Welty
1910 Oakley .Philip Douglas
1910 Pratt, Ishmael Charles
1910 Sabastian, Thiruchelvam
1910 Shaw, Hugh Thomas
1910 Sieger, Edward Louis
1910 Sousa, Pascal John de
1910 Souza, Antonio Bernardo de
1910 Waterhouse, John Howard
1910 White, Maurice Forbes
1911 Blacklock, Breadalbane
1911 Brown, Frederick Forrest
1911 Chand, Diwan Jai
1911 Holmes, John Morgan
1911 levers, Charles Langley
1911 lies, Charles Cochrane
1911 Ingram, Alexander
1911 Kirkwood, Thomas
1911 Knowles, Benjamin
1911 Lid die, George Marcus Berkeley
1911 Lomas, Emanuel Kenworthy
1911 Mackarell, William Wright
1911 MacKnight, Dundas Simpson
1911 Mascarenhas, Joseph Victor
1911 Murray, Ronald Roderick
1911 Oluwole, Alddiya Ladapo
1911 Rao, Koka Ahobala
1911 Sinton, John Alexander
19x1 Tarapurvalla, Byramji Shavakshah
1911 Taylor, John Archibald
1911 Woods, William Medlicott
191a Acria, Joseph Reginald
1912 Anderson, Edmund Litchfield
191a Borle, James
191a Bowie, John Tait
Data of
Diploma
1 9f a Braasey, Laurence Perdval
1912 Christie, David
19x2 Dillon, Henry de Courcy
1912 Dunn, Lillie Eleanor
19x2 Hardwicke, Charles
1912 Jagose, Jamshed Rustomji
1912 Kochhar, Mela Ram
1912 McGusty, Victor William Tighe
1912 Milne, Arthur James
1912 Mitra, Manmatha Nath
1912 Myles, Charles Duncan
1912 Felly, Huntly Nevins
1912 Prasad, Bindeshwari
1912 Prentice, George
1912 Ross, Frank
1912 Russell, Alexander James Hutchison
1912 Ruthven, Morton Wood
1912 San dilands, John
1912 Seddon, Harold
1912 Smalley, James
1912 Strickland, Percy Charles Hutchison
1912 Watson, William Russel
1913 Austin, Charles Miller
1913 Banker, Shiavux Sorabji
1913 Becker, Johann Gerhardus
1913 Carrasco, Milton
1913 Clark, James McKillican
1913 Forsyth, Charles
1913 Grahame, Malcolm Claude Russell
1913 Grieve, Kelbume King
19x3 Hargreaves, Alfred Ridley
1913 Hepper, Evelyn Charles
1913 Hiranand, Pandit
1913 Jackson, Oswald Egbert
X913 Khaw, Ignatius Oo Kek
19x3 MacKelvie, Maxwell
1913 MacKinnon, John MacPhail
1913 Macmillan, Robert James Alan
1913 Mouat-Biggs, Charles Edward Forbes
1913 Noronha, John Carmel
1913 O'Connor, Edward
1913 Olubomi-Beckley, Emanuel
1913 Pestonji, Ardeshir Behrxmshah
1913 Puttanna, Dodballapur Sivappa
1913 Reford, John Hope
19x3 Smith, Edward Arthur
X913 Stewart, Samuel Dudley
1913 Walker, Frederick Dearden
19x3 Wilbe, Ernest Edward
1913 Wilson, Hubert Francis
1913 Yin, Ulg Ba
1913 Young, William Alexander
1914 Arculli, Hassan el
1914 Chohan, Noormahomed Kasembha
1914 Connell, Harry Bertram
19x4 Gerrard, Herbert Shaw
1914 Gimi, Hirji Dorabji
19x4 Gwynne, Joseph Robert
1914 Hodkinson, Samuel Paterson
1914 Jackson, Arthur Ivan
1914 Kaushash, Ram Chander
1914 Kelsall, Charles
19x4 Luanco y Cuenca, Maximino
1914 Misbah, Abdul-Ghani Naguib
Data of
Diploma
1914
Nasdu, Bangalore PasupuUti
Balakrishna
[914 Rowe, John Joseph Stephen
[914 Roy, Raghu Nath
914 Shiveshwarkar, Ramchandra Vishnu
914 Sur, Sachindra Nath
914 Talati, Dadabhai Cursedji
914 Wilkinson, Arthur Geden
914 Wright, Ernest Jenner
(91$ Lobo, John Francis
[915 Madhok, Gopal Dass
[915 Pearson, George Howorth
1915 Swami, Karumuri Virabhadra
915 Wood, John
>916 Barseghian, Mesroob
[916 Chaliha, Lakshmi Prasad
[916 Lim, Albert Liat Juay
1916 Lim, Harold Liat Hin
[916 Metsger, George Nathaniel
[9x6 SdderstrGm, Enk Daniel
[916 Wheeler, Louis
1917 Chapman, Herbert Owen
1917 Krishnamoorthy, Yedatore Venkoba
1917 Lipldn, Isaac Jacob
1918 Watts, Rattan Claud
1919 Bowie-Evans, Charles Harford
19x9 Bumie, Robert McColl
1919 Celestin, Louis Abel
X919 Cummings, Eustace Henry Taylor
1919 Darling, Georgina Renington
X919 Drake, Joan Margaret Fraser
X919 Fraser, William James
1919 Gordon, Rupert Montgomery
X919 Krige, Christian Frederick
X919 Maplestone, Philip Alan
19x9 Oluwole, Isaac Ladipo
19x9 Rustomjee, Khusshuyee Jametidjee
1919 Sawers, William Campbell
1919 Thompson, Mary Georgina
1919 Turner, Gladys Maude
1919 Young, Charles James
1920 Adler, Saul
1920 Anderson, William Jenkins Webb
1920 Campbell, George
X920 Cobb, Charles Eric
1920 Cobb, Enid Margaret Mary
1920 Fernandez, Daniel David
1920 Lim, Chong Eang
1920 McHutcheson, George Browne
1920 van der Merwe, Frederick
1920 O'Farrell, Patrick Theodore Joseph
1920 Renner, Edo wo Awunor
1900 Vaughan, James Church will
1920 WalleT, Harold William Leslie
1921 Nixon, Robert
1921 Richmond, Arthur Stanley
1921 Skinner, James Macgregor
1921 Stewart, Robert Bell
1921 Thomson, Marion
ANNALS OF TROPICAL MEDICINE
AND PARASITOLOGY
EDITORIAL NOTICE
Articles for publication should not exceed twenty-five pages of
the Annals, and will be understood to be offered alone to this
Journal. They should be typewritten and addressed to:—The
Editors, School of Tropical Medicine, The University, Liverpool.
Illustrations for text figures or charts should be drawn clearly
and firmly in Indian ink, if possible on Bristol board. N.B .—Blue
or ‘other coloured ruling in squares or lines cannot be reproduced.
All lettering, names or legends on text-figures, charts or maps
should be printed sufficiently large to allow of clear legibility on
reduction if necessary.
Plates and illustrations should be accompanied by short explana¬
tions.
References to authors in the text must be made in the following
way:—‘According to Smith (1900) the spleen is enlarged, but
Robinson (1914) says the reverse.' The references should be
collected in alphabetical order of authors' surnames at the end of
the paper, and arranged in the following way: —
Robinson, S. (1914). 4 The spleen in malaria.* Annals of Nosology ,
Vol. XX, pp. 20-25.
Smith, T. (1900). 4 Enlargement of the spleen in malaria.* Journal of
Patnometry , Vol. I, pp. 1-20.
Fifty reprints are supplied to each author, free of charge.
Additional copies (up to 200) can be supplied at cost price.
Subscription: £1 2s. 6d. per volume, post free, payable in
advance to the Secretary, University Press, 57 Ashton Street,
Liverpool, to whom correspondence concerning advertisements
should also be addressed.
LIVERPOOL SCHOOL OF TROPICAL
MEDICINE
SCIENTIFIC. RECORD
COMPILED BY
DOROTHY ALLMAND
Librarian of the Liverpool School of Tropical Medicine
Plates I—IX
The new laboratories of the Liverpool School of Tropical
Medicine were formally opened on 24th July, 1920, by Lord
Leverhulme, Honorary Vice-President and former Chairman of the
School.
The building had been placed, in 1915, at the disposal of the
Military authorities for the purposes of a hospital, and was used
by them until 1919, for the treatment of patients suffering from
tropical diseases, mainly malaria and dysentery. The hospital,
which consisted of about 200 beds, was under the charge of
Lieut.-Col. J. W. W. Stephens, R.A.M.C., Professor of Tropical
Medicine, and during its four years’ existence more than 3,000 cases
were admitted.
Subsequent to evacuation by the Military authorities, the
decoration and equipment of the laboratories were put in hand, and
to-day the School affords excellent conditions and facilities for
instruction and research.
The laboratories are situated in the University grounds, and
dose to the Royal Infirmary, in which the School has its Tropical
Ward and adjoining clinical laboratory. In addition to the
basement, which contains the Photographic Department and large
storage rooms, there are four floors. The ground floor comprises
the Lecture Theatre, with accommodation for about seventy
students; the Library, where, besides text-books and miscellaneous
scientific publications, over one hundred current medical journals
from all parts of the world can be consulted; and the Museum, a
spacious room, 80 feet by 69 feet, with preparation room adjoining.
2
On the first floor are the Departments of Tropical Medicine and
Entomology, the latter with its library of specialised literature, and
six research rooms. The second floor comprises the excellently
lighted class laboratory, 69 feet by 58 feet, and four research rooms
devoted to the Department of Parasitology, while the third floor
has a large research laboratory .and two smaller research rooms. On
the roof are an Insectarium, a mosquito-proofed house, and the
animal houses.
The occasion of the .official opening was marked by the issue
of a volume* which traces the history of the School from its
foundation down to the year 1920. The main purpose of this
publication was to perpetuate the names of those who have been
closely associated with the School in its varying activities in the
past, and this aspect may here be briefly summarised before
proceeding to an account of its scientific activities which, in the
‘ Historical Record,’ are merely outlined.
The Liverpool School of Tropical Medicine was founded in 1898
by the late Sir Alfred Lewis Jones, K.C.M.G., a prominent
Liverpool ship-owner, who, fired by an appeal for the study
of tropical diseases issued by the late Rt. Hon. Joseph Chamberlain,
then Secretary of State for the Colonies, offered* to the President of
the Royal Southern Hospital the sum of £350 for three years, to
promote the special study of such diseases. Quarters were obtained
in the Pathological Department at the University, and a Dean was
appointed in the person of Professor Rubert Boyce, to whose tireless
energy much of the early success of the School was due. In
February, 1899, Dr. H. E. Annett was appointed Demonstrator in
Tropical Pathology; in April, Major Ronald Ross accepted the post
of Lecturer, and the School was officially opened; and in May
teaching commenced.
The early history of the Liverpool School is a record of struggle;
it was not founded by the Government, as was the London School;
it had no grant, no assured income, nor even, at first, Government
recognition. The School early turned its attention to research, and
in the summer of 1899 the first of its expeditions was despatched
to Sierra Leone. During the next fourteen years funds were
# Liverpool School of Tropical Medicine: Historical Record, 1898-1920. At the
University Press > 1920. viii, 103 pp., 36 plates. 10/6 post free.
3
collected to equip and maintain more than thirty expeditions to
various tropical regions, including West and Central Africa, Brazil
and the West Indies.
In the meanwhile, at home the School was enlarging its scope
and consolidating its position. In 1900 the Government gave it full
recognition by placing it on the same terms as the London School
with regard to newly-appointed medical officers and their courses
of training, and four years later further recognition was received in
the form of a financial grant from the Colonial Office, which grant
was later increased and has continued to the present day. In 1901
a Lectureship and Fellowship were founded, in commemoration of
the work of the late Walter Myers, who died of yellow fever in
Brazil while serving with the Fourth Expedition. Major Ross was
appointed to the former, and the latter was filled by Dr. J. E.
Dutton, who, four years later, died in the Congo while investigating
tick fever. In 1903 the Alfred Jones Chair of Tropical Medicine
was founded, the first holder being Major Ross, while Dr. J. W. W.
Stephens, who had joined the staff in the previous year, was
appointed to the vacant Walter Myers Lectureship. In 1905
another important Lectureship was founded, namely, that of
Economic Entomology, which was accepted by Mr. R. Newstead,
of Chester. As the work and influence of the School increased,
many new appointments were made, so that now, at the age of
twenty-two, the School has three University Chairs,* namely,
Tropical Medicine, Entomology and Parasitology, filled by
Professors Stephens, Newstead and Yorke, respectively, assisted by
a staff of eight Lecturers.
In 1903 the School moved into new laboratories, and in the
following year a Diploma in Tropical Medicine was instituted.
In 1904 the Runcorn Research Laboratories were established for
the purpose of conducting investigations on the large collection of
trypanosomes and other protozoa, which had been amassed by
members of the expeditions on various occasions. These labora¬
tories were first placed in the charge of Dr. Wolferstan Thomas, and
during their ten years’ existence much valuable research was done
there.
• A fourth Chair, that of Tropical Diseases of Africa, has now (1921) been established,
and is held by Professor B. Blacklock.
4
In 1906 was issued the first number of the Annals of Tropical
Medicine and Parasitology . This publication had been preceded
by a series of twenty-one Memoirs , mostly reports of the School's
expeditions.
In 1911 the Yellow Fever Bureau came into being with
Dr. Harald Seidelin as Director. As the name implies, the Bureau
was established for the purpose of promoting the study of the many
problems surrounding this disease. A journal was issued, the
Yellow Fever Bulletin , of which three volumes were published.
Clinical instruction to students of the School had formerly been
given in a special ward at the Royal Southern Hospital. This
hospital being at some distance from the laboratory, arrangements
were subsequently made tfith the Royal Infirmary for the erection
of a new tropical ward in the Infirmary grounds. The ward, which
contains ten beds, with an adjoining clinical laboratory, was opened
in 1914.
On the arrival of the Fifteenth (Yellow Fever) Expedition at
Manaos in 1905, it was found necessary for the work of the
Expedition to establish a laboratory of a more or less permanent
character. The Mandos Research Laboratory remained in existence
until January, 1909, when, owing to the return of its Director,
Dr. Wolferstan Thomas, to Liverpool, it was closed. In June,
1910, Dr. Thomas returned to Mandos and opened the present
laboratory. In 1914 it was decided to extend, the activities of this
branch of the School; the outbreak of war, however, caused all
developments to be deferred until 1919, when three research
assistants were appointed to the laboratory.
With a view to carrying oil research work in tropical medicine,
the School desired to establish a permanent laboratory on the West
Coast of Africa. Funds were available, through the munificence
of the late Sir Alfred Jones, and a suitable site on Tower Hill,
Freetown, Sierra Leone, having been placed at the disposal of the
School by the Colonial and War Offices,, the laboratory is now in
course of erection.
Annals Prop. Med. & Parasitol., Vol. AT
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5
The following brief record of the research work of the School is
dealt with under subjects; no attempt at a critical estimate has
been made.
MALARIA AND SANITATION
The first Expedition left England in July, 1899, for Sierra
Leone, and consisted of Major Ross, Dr. H. E. Annett, Dr. R.
Fielding-Ould, Mr. E. E. Austen, officially appointed by the
British Museum, and Dr. Van Neck, delegate of the Belgian
Government to the School. The purpose of the Expedition was to
study malaria in man, and a report of the work accomplished was
published in 1902 as Memoir II. The investigators discovered two
species of Anopheles in Freetown, namely, A. funestus , Giles, and
A . costalis, Loew, and by dissection established the presence of
blasts (sporozoites) in them. They concluded that both A. costalls
and A. funestus are hospitable to the human Haemamoebidae; that
A. co stalls is hospitable to all three of the human species, and
A. funestus certainly to H. malariae t and probably H . vivax ; no
observations were made regarding the connection of A. funestus
with H. praecox . They further pointed out the difference between
Culex and Anopheles , and studied more especially the bionomics of
the latter. 1 Precautions against the bites of gnats * and ‘ operations
for reducing the number of Anopheles' were fully considered. Dr.
Fielding-Ould continued the work of the Expedition at Freetown
after the other members had returned to England, and he also
prosecuted his researches at Accra and Lagos, his report of which
is included in Memoir II. At Freetown six of twenty-nine
Anopheles dissected showed parasites. Seventeen Culex gave
negative results. At Accra fifty-two Culex were negative. At
Lagos thirty-seven Anopheles were dissected and zygotes found in
one. The reports dealt mostly with the sanitary survey of the
districts.
In 1900 a third Malaria Expedition was sent to Northern and
Southern Nigeria. This Expedition, which was composed of
Drs. H. E. Annett, J. E. Dutton and J. H. Elliott, spent six months
in Nigeria, and in Memoirs III and IV reports were published of the
work done, the former relating to malaria and the latter to filariasis
(see p. 37). Seven of two hundred and eighty-one Anopheles
6
dissected at Bonny were found to be infected with malaria; it was
noted that it was difficult to decide what was the exact type of
parasite present. The observation made by Koch and by the
Royal Society’s Commission that native children are infected with
malarial parasites to a large degree was confirmed; the incidence of
infection is given in the following table. It was also proved that
Tabu showing the total number 7 of’children'examined and found infected
throughout Nigeria.
1
Ages
Number examined
Number infected
Percentage infected
0-5
220
U 4
5 r8
5 -io
1 108
2 7
25*0
10 +
40
4
10*0
quartan and simple tertian parasites exist in West Africa, a fact
contrary to the experience of the Royal Society's Commission of
the previous year. The bionomics of Anopheles were studied still
further and a series of experiments in propagation carried out,
confirming and elaborating the discovery of the previous Expedition
that the female mosquito requires a meal of blood both for fertilisa¬
tion and for the development of the ova. In their recommendations
concerning prophylaxis they advocated (i) the segregation of
Europeans at a distance of about half a mile (a principle already
put forward by the Royal Society's Commission on Malaria also at
work in West Africa), and (2) the surface drainage of areas around
their quarters. In an appendix they gave charts and descriptions
of cases of hyperpyrexial fever, first described by Thompstone and
Bennett. The hyperpyrexial stage lasts one to three weeks, and is
followed by very extended lysis. An exhaustive account was given
in the second report ( Memoir IV) of the mouth parts of the female
A. costalis , and, in an appendix by Theobald twenty-five mosquitoes
were described of which nine were new species, namely, Stegomyia
irritans , S. nigricephala i Culex duttoni , C. decens , C. pruina ,
C. invenustus , C. nebulosus, C. rima and C. invidiosus (with
figures).
In June, 1901, Major Ross and Dr. Logan Taylor went to
Freetown to organise an anti-mosquito campaign. Work was
7
commenced by engaging the services of between thirty and forty
men, who were divided into two gangs: the Culex gang and the
Anopheles gang. The former collected from private houses all the
broken bottles and buckets, empty tins, etc., in which Stegotnyia
and Culex breed; the latter drained the pools and puddles in which
Anopheles breed in the streets and backyards of the houses.
Reports cm the progress of this campaign were published as
Memoir V, Parts i and 2, and later Ross issued a small book:
‘ Mosquito Brigades, arid how to organise them.’
In September, 1901, Dr. Dutton went to the Gambia and
inspected the conditions of health there, with the result that the
Colony organised measures similar to those in operation at Freetown
(Memoir X). About 32 per cent, of the infections examined
Table ihowing the total number of children examined and found infected
throughout the Gambia.
Ages
Number examined
Number infected
Percentage infected
°*5
78
64
82*0
5-10
22
20
91*0
10-f
>3
7
53-8
showed quartan parasites, and about a third of the malignant
tertian cases showed crescents. The simple tertian parasite was
found three times only.
Dutton noted the universal infection of canaries with
Halteridiunt, which was also found in other birds.
Of twenty-four A. funestus dissected, one contained sporozoites
and one zygotes.
A . costalls was found breeding in boats, street drains, wells, tubs
and barrels, and in tidal water containing 1*7 per cent, salt,
together with C. thalassius. It would appear that A. funestus and
its varieties are rural mosquitoes, while A. costalis is essentially
town bred and capable of utilising any small collection of water for
breeding purposes.
In an appendix, descriptions of eighteen species were given by
Theobald, including A. costalis, var. melas; A. funestus , var.
8
umbrosus\ A. funestus , var. subumbrosus\ Stegomyia albocephala ,
sp. n.; Culex annulioris , var. gambiensis , v. n.; C. anarmosttfs ,
sp. n.; £ 7 . thalassius , sp. n. ; C. eucldstus , sp. n. ; Lasioconops
poicilipes , gen. n., sp. n.; Corethra ceratopogones , sp. n. Eight
specimens of 6. longipalpis var. tachinoides were taken. Dutton
recorded the finding of a filarial embryo in the thoracic muscle of
£ 7 . anarmostus.
It was during his researches into the blood parasites of the
Gambia that Dr. Dutton made a discovery of the highest scientific
importance, namely, the identification for the first time of a trypano¬
some in the blood of a man, a patient of Dr. Forde. This parasite
was subsequently (see p. 16) shown to be the cause of sleeping sickness,
and was named Trypanosoma gambiense , Dutton, 1902.
In 1901, Dr. C. Balfour Stewart was sent to the Gold Coast, to
conduct operations similar to those carried on at Sierra Leone, while
early in 1902 Major Ross again visited Freetown to ascertain
by a thorough inspection the results of the previous Expedition.
In the autumn of this year, Dr. Logan Taylor proceeded to Cape
Coast Castle, Gold Coast, and reported upon the sanitary conditions
prevailing there, with suggestions as to their improvement.
(Memoir VIII).
In September, 1902, by request of the Suez Canal Company,
Major Ross went to Ismailia to investigate the causes of the
prevalence there of malaria, and to recommend measures for its
prevention. Anopheles and Culex were found breeding in water
containing 0*9 per cent, of salt. He concluded that the majority of
Anopheles which caused malaria in Ismailia came from the marshes
in immediate proximity to the town. Prophylactic measures based
on his recommendations were commenced immediately ( Memoir IX).
In February, 1904, sixteen months later, Professor Boyce visited
Ismailia; from statistics furnished in his report {Memoir XII), it
appeared that the number of cases of malaria had fallen from 1,551
in 1902, to 209 in 1903.
During 1904, two Sanitary Expeditions were despatched to West
Africa: one to Bathurst, Conakry and Freetown, and the other to
the Gold Coast. Reports were published in 1905 as Memoirs XIV
and XV.
In May, 1906, Professor Ross went to Greece in order to advise
Annals Trop. Med, £? ParasitolVol, XV
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SECOND LABORATORY - OF THE SCHOOL
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9
the Lake Copais Co. with regard to anti-malaria measures. In the
following year, at the request of the Colonial Office, he visited
Mauritius for the same purpose; his report, entitled ‘ The Prevention
of Malaria in Mauritius,’ was published in 1908 (Waterlow & Sons).
In 1908, the Twenty-first Expedition of the School, consisting
of Professor Newstead, Dr. W. T. Prout and Dr. Alan Hanley, was
despatched to Jamaica. Reports were published in the Annals of
Tropical Medicine and Parasitology, Vol. Ill, pp. 421 and 471, the
first, dealing with medical and economic entomology, by Professor
Newstead (see p. 39), the second, on malaria, by Dr. Prout. From
data furnished by Dr. Neish, of Spanish Town, the frequency of
the various species of malaria parasite was found to be approxi¬
mately : simple tertian 54 per cent., malignant tertian 36 per cent.,
quartan about 1 per cent., and mixed infections about 8 per cent.
The occurrence of several cases of blackwater fever was noted. The
enlarged spleen rate in different parishes varied from o to 60 per
cent. Prout found that the average percentage of malarial deaths
to total deaths was nearly 20 per cent., that in four years the total
admissions to hospitals from malaria had increased by 55 per cent.,
and that over 33 per cent, of the total admissions were due to
malaria; it was reported that the annual cost of treating malarial
patients' was over £6,000. Various anti-malarial measures were
recommended. In 1909, Boyce visited Jamaica, and reported upon
the work done by the Commission appointed by the Governor in
October, 1909, to deal with the malaria problem {Annals, Vol. IV,
p. 233). It was recorded that one-half of the island, namely, those
parts above 1,000 feet, was practically free from malaria-carrying
mosquitoes.
Ross and D. Thomson, working in Liverpool, published a paper
entitled *Sorfie Enumerative Studies oh Malarial Fever’ ( Proc.
Roy. Soc., B., Vol. LXXXIII, p. 159, and Annals, Vol. IV,
p. 267). Some of the conclusions were: (1) No fever exists unless
the parasites exceed from 500 to 1,500 per c.mm.; (2) the parasites
tend to remain continuously in the blood in small numbers between
the febrile relapses; (3) close correlation between the number of
parasites and the amount of fever caused by them; (4) studies on
quinine gave a numerical estimate of its effect, a few days’ use of
the drug reducing the parasites by from 50 per cent, to 80 per cent.;
IO
(5) crescents apparently require eight to ten days for development;
quinine affects their numbers only by destroying the generating
cells. Thomson, who subsequently investigated the life history of
crescents {Annals, Vol. V, p. 57), concluded that these do not live
for more than a few days in the peripheral blood. They are
replenished from surviving asexual forms, and quinine has no
action on crescents but only on the asexual source of supply. He
concluded {Annals, Vol. VI. p. 223) that administration of quinine in
doses of 20 grains daily for three weeks is almost certain to destroy
both the asexual and sexual parasites. In a study of the leucocytes
in malarial fever (ibid., p. 83), he stated that malaria could be
diagnosed by the leucocytic formula.
In 1912, J. G. Thomson and McLellan confirmed Bass’s observa¬
tions on the cultivation of malarial parasites. In twenty-four hours
P. falciparum was found to undergo segmentation, the maximum
number of merozoites counted being thirty {Annals, Vol. VI,
p. 449). In the case of P. vivax {Annals, Vol. VII, p. 153), sixteen
merozoites were produced. These cultures of P. vivax differed from
those of P. falciparum in that there was no tendency to clumping.
Further experiments in the cultivation of P. falciparum and
P. vivax were made {ibid., p. 509), when it was noted that the
optimum temperature for cultures was 38° C.
Sinton, investigating Uriola’s test of malarial pigment in the
urine {Annals, Vol. VI, p. 376), concluded that it is almost
impossible to exclude extraneous pigment.
In 1912, Dr. David Thomson was sent to Panama to study
certain malarial problems with Dr. James, Chief Assistant
Physician to the Ancon Hospital. The first part of his report
{Annals, Vol. VII, p. 125) dealt with the sanitation in the Canal
Zone, Trinidad and British Guiana; the second {Annals, Vol. VIII,
p. 85) with the origin and development of gametes in malignant
tertian malaria. He noted that they develop chiefly in the bone
marrow and in the spleen, the period of incubation being about ten
days.
While examining a malarial blood film sent from the Central
Provinces of India, Stephens was struck by the peculiar appearance
of the parasite {Proc. Roy. Soc., B. Vol. LXXXVII, p. 375, and
Annals, Vol. VIII, p. 119). It exhibited the following peculiarities:
II
(i) it was extremely amoeboid; (2) the cytoplasm was scanty; (3) the
nuclear protoplasm was out of proportion to the volume of the
parasite. It differed from P. falciparum by its amoeboid activity,
and from P. vivax by its smaller size, the delicate nature of its
amoeboid processes, the irregularity of its chromatin and the rarity
of typical ring forms. Stephens proposed to name this parasite
P. tenue. Later he described peculiar forms of a malaria parasite
in a blood slide from the Gold Coast {Annals, Vol. IX, p. 169).
In addition to these, large and apparently quite normal quartan
parasites occurred, and it was possible to trace a transmission from
normal ring forms to those in which chromatin particles or strands
without any protoplasm were seen in the red cells.
In 1917, the School was asked by the War Office to undertake
investigations into the treatment of malaria; the results of this work
were published in thirty papers {Annals, Vols. XI-XIII).
No case was considered to be malaria unless parasites were
found. The results of treatment were in all cases controlled by
daily microscopical examinations combined with the clinical record.
Simple Tertian. The investigators established: (1) that intra¬
muscular injection of quinine bihydrochloride was an effective
method of treating a malarial attack, a matter which had been one
of considerable dispute in the medical press just previous to this
work. (2) That for the palliative treatment of malaria, that is, for
keeping a person free from relapses over long periods, it was better
to give a certain total amount of quinine on each of two consecutive
days than on each of six days in the week: thus, 60 grains
administered as 30 grains for two days gives a better result than
the same amount administered as 10 grains for six days. (3) The
best palliative result was obtained by administration of 45 grains
on each of two consecutive days weekly over a period of two
months. (4) They found that a certain treatment may give a
certain ‘ curative ’* result on one occasion, while the same treatment
repeated on another occasion might give a quite different result.
(5) Novarsenobillon was found to be as efficacious as quinine in the
treatment of paroxysms, but its curative effect, like that of quinine,
was practically nil. (6) The best ‘curative* result was'obtained by
• The term cure was used to signify no relapse within an observation period of 66 days
after cessation of treatment.
12
the administration of Liquor arsenicalis, minims 30, daily over a
period of eight weeks in combination with two initial intramuscular
injections of quinine bihydrochloride, grains 15, on two days only.
Owing to the Armistice, these observations were not repeated.
The intravenous injection of quinine was found to have no real
curative effect.
The relapse period after treatment of eight hundred simple
tertian cases was recorded {Annals, Vol. XIII, p. 125). It is
essential to recognise that these figures were based on an observation
period of sixty days. The incidence is shown in the accompanying
graphs. It will be seen that of those cases that relapse the majority
do so in the first twenty days after cessation of treatment. Further,
the time of occurrence of the paroxysms was noted in one thousand
cases {Annals, Vol. XIV, p. 365). The majority occurred at^
2 p.m. with the conditions of life under which the patients (soldiers)
were living; over 90 per cent, occurred during the hours of activity,
that is, between the hours of 7 a.m. and 6.59 p.m.
Malignant Tertian. Neither a single. nor a series of six
intravenous injections of quinine bihydrochloride (grains 10 to 15)
caused the disappearance of parasites, either trophozoites or
gametes, from the peripheral blood, whereas in the case of simple
tertian the disappearance was rapid. Under quinine treatment,
grains 30 to 45 daily, crescents did not persist in the peripheral
blood in the majority of cases for more than three weeks. How long
they persist without quinine was not determined.
In July, 1919, Blacklock and Carter recorded the experimental
infection, for the first time, of Anopheles plumbeus with P. vivax.
Experiments were made {Annals, Vol. XIII, pp. 187 and 413), with
the result that the observers were able to obtain infections of the
gut and salivary glands of laboratory-bred A. plumbeus at a
temperature of 28° C.; at room temperature gut infection only was
obtained. Infection of the gut was also produced with P. vivax in
the case of A. bifurcatus at 28° C. Later experiments {A nnals,
Vol. XIV, p. 275) with A. plumbeus resulted in gut infection with
oocysts of P. falciparum .at 28° C.
•3
Graph i.
Percentage of total relapses
in each 20-day period.
Graph 2.
Percentage of cases treated
which relapse in..
each 20-day period.
Graph 3.
Percentage of cases treated
nofr haring previously
relapsed which do so in
each 20-day period.
Days after cessation of treatment.
*4
BLACKWATER FEVER
In 1907, the Nineteenth Expedition, consisting of Dr. J. O.
Wakelin Barratt and Dr. Warrington Yorke, was despatched to
Nyasaland to study blackwater fever, a report being subsequently
published in Annals , Vol. Ill, p. 1. The object of the investigators
was to trace out some of the internal processes, the terminal event
of which is the appearance of blackwater, believing that in that way
many obscure points in connection with the causation and treatment
of the condition would be cleared up. The first point was to
determine the action of quinine, acid and alkali upon the red
cells during blackwater fever. They found that haemolysins,
present in the blood, played no part in the production of black¬
water. It was considered that the suppression of urine is due to
a mechanical blocking of the renal tubes by the formation of large,
firm, coarsely granular casts in the ducts of Bellini. In a later
study {Annals, Vol. V, p. 287), on the suppression of urine in black¬
water fever, Yorke and Nauss re-investigated the mechanical theory
and found that it is considerably facilitated by any factor which
tends to lower the blood-pressure, and by that means the secretion
of water by the glomeruli, but that if the blood-pressure is kept up
by the injection of saline solutions, the tendency to suppression is
decreased. Arising out of these latter experiments, the passage of
haemoglobin through the kidneys was studied by Yorke (ibid.,
p. 401), who was led to consider that haemoglobin is excreted by
the renal epithelium rather than filtered through the glomeruli, and
that the amount of haemoglobin eliminated into the urine is
dependent upon the activity of the epithelium lining the renal
tubules.
It was shown by Simpson ( Annals, Vol. VI, p. 313) that the
haemoglobin liberated from the red cells in malaria escapes in larger
quantities by the faeces than by the urine. The study of haemo¬
globin metabolism in blackwater fever was continued, and a report
made (Bio-Chemical Journal, Vol. V, p. 378) on the quantitative
estimation of urobilin in the excreta. In later observations on
haemolysis in malaria (Annals, Vol. VI, p. 231), Simpson concluded
that the serum of malarial patients may possess the power of
haemolysing normal red blood cells. The haemolytic effect could
Annals Trap. Med, Parasitol,, J'ol, XV
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FRONTAGE OF THIRD LABORATORY OF SCHOOL
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not be obtained at all periods of the paroxysm, nor in every case;
it appeared to be produced at the period of sporulation, and rapidly
disappeared. Simpson and Edie (ibid., p. 443), observing the
excretion of urobilin in animals and man, found that an increase
may occur after the administration of quinine in doses of 10 to 30
grains a day, and that a similar result follows injection of blood
pigment or haemolytic drugs.
Experiments were devised by Barratt and Yorke ( Annals ,
Vol. VIII, p. 509) for examining the relation of bile pigments to
haemoglobin. Experimenting with rabbits, they found that
consequent upon intravenous injection of haemoglobin solution there
was a distinct and immediate increase not only in the concentration
of the bile pigment, but also in the amount of bile pigment excreted.
Two hypotheses were advanced to explain this increase: (1) that
the haemoglobin injected is actually converted by the liver into bile
pigment; or (2) that it merely stimulates the liver cells to an
increased production of bile pigment.
Stephens ( Thomfson-Yates Lab. Reports , Vol. V, Pt. 1, p. 193)
recorded that blackwater fever occurred in eleven of twenty-two of
the United States. An account was given of the distribution of the
disease, and an extensive bibliography appended. An analysis of
ninety-five cases showed that when the blood was examined before
the onset of blackwater, malarial parasites were present in 95*6 per
cent, of cases, whereas on the following day the remarkable fall to
17*1 per cent, was the result. On the day of blackwater itself the
figure was 6 i'g per cent.
A series of studies in blackwater fever were made by Stephens
(Annals, Vol. VII, p. 479). The subject was considered under the
following headings: (1) malarial parasites; (2) pigmented leuco¬
cytes; (3) post-mortem examinations; (4) influence of malaria;
(5) relationship to species of malaria parasites; (6) effect of period of
residence; (7) seasonal prevalence; (8) correlation between malaria
and blackwater statistics; (9) second attacks. A schedule for
recording cases of blackwater was devised and recommended
(Annals, Vol. VIII, p. 639). The results were recorded (Annals,
Vol. IX, p. 201) of a statistical examination of the respective times
at which quinine was given and blackwater occurred. Graphs were
published showing that correlation existed between the time of
i6
taking quinine and the onset of symptoms, but this did not
necessarily imply any relationship of cause and effect. Data on the
duration of haemoglobinuria were collected ( Annals, Vol. IX,
p. 539). In one hundred and sixty-seven records it was found that
the duration was not more than twelve hours in a quarter of the
cases, not more than one day in half the cases, and not more than
two days in three-quarters of the cases. Finally, the importance of
furnishing population statistics in connexion with cases of black-
water fever was emphasised ( Annals, Vol. X, p. 345).
PIROPLASMOSIS
At Runcorn, Brcinl and Hindle studied the morphology of Piro-
plasma cants (Annals , Vol. II, p. 233), and worked out the nuclear
details of the parasite. Later, Breinl and Annett (Hid., p. 383)
concluded that the haemolysis in Ptroplasma cants is not due to a
formation of a specific haemolysisin or isolysin, but to the mechanical
disintegration of the red blood corpuscles after the escape of the
parasites from them. Barratt and Yorke found that in piro-
plasmosis the haemoglobinuria was attended with and dependent
upon haemoglobinaemia. Of one hundred and forty-three cattle
examined at Sierra Leone {Annals, Vol. IX, p. 418) about 5 per
cent, were found to be infected with P. bigemtnum, while Theileria
mutans was encountered in between 20 and 30 per cent.
TRYPANOSOMIASIS
In view of the great importance of the discovery of the trypano¬
some in man by Dr. Dutton* in 1901, described and named by him
77 gambiense (Thompson-Yates Reports, Vol. IV, p. 455), an
Expedition was sent out to the Gambia and French Senegal
in September, 1902, in charge of Dr. Dutton and Dr. J. L. Todd,
for the study of Trypanosomiasis. While these investigators were
at work abroad, the first case, that of a European who had returned
with Dr. Dutton in 1901, remained in Liverpool, and Dr. Annett
infected monkeys and 25 per cent, of tame rats successfully, but did
not succeed in infecting tame mice, rabbits or guinea-pigs. One of
the infected monkeys died, but the other recovered, and no parasites
• For the history of the discovery see Boyce, Ross and Sherrington, Lancet , Feb. 21, 1903.
could be found by sub-inoculation into rats. The results of his work
were incorporated in the report of the Expedition (Memoir XI). On
reaching the Gambia, one thousand and forty-three natives were
examined, the majority of whom were children or young adults, and
apparently healthy; six were found to be infected. Trypanosomes
were also found in the blood of a quadroon. Clinical descriptions
were furnished in the report of these first eight cases of trypano¬
somiasis, of which those of the European and the quadroon
terminated fatally after a duration of about eighteen months. Of
thirty-six horses examined, ten were found to be infected with
trypanosomes. Transmission experiments were made with Glossina
palpalis and later with Stomoxys, but with negative results. A
series of inoculations were made of the human and equine trypano¬
somes in experimental animals, the results of which led Dutton and
Todd to the conclusion that the parasites were not of the sa m e
species; the Gambian horse trypanosome was subsequently named
T. dimorphon. Several new species of flagellates occurring in
birds, mice, tortoises, etc., were described in this report, including
T. johnstoni, T. mega and T. karyzeukton.
Prior to the return of this Expedition, the discovery of trypano¬
somes in the cerebro-spinal fluid of cases of sleeping sickness in
Uganda by members of the Sleeping Sickness Commission of the
Royal Society, caused the subject of trypanosomiasis to assume great
importance. At the invitation of King Leopold, an Expedition was
sent in 1903 to study sleeping sickness in the Congo Free State,
consisting of Drs. Dutton, Todd and Christy. The results of these
investigations were incorporated in Memoir XIII, and illustrated the
occurrence and distribution of trypanosomiasis, described the
symptoms of the disease in all its stages, both in Europeans and'
natives, and showed how sleeping sickness, so-called, is related to
trypanosomiasis as a symptom of that disease.
They first stated that they "were unable to find any difference
between the trypanosome occurring in cases of sleeping sickness in
the Congo and T. gambiense. There was a very evident clinical
connection between cases with only very slight symptoms (trypano¬
soma fever) and advanced cases of ‘ sleeping sickness.’ In twenty-
five of thirty-eight cases they found parasites in the cerebro-spinal
fluid, adopting Quincke’s new method of diagnosis by lumbar
puncture. They infected rats, mice, rabbits, guinea-pigs, and
studied the morphology of the Congo and Gambian trypanosome in
these animals. They noted that about 50 per cent, of such inocula¬
tions failed, and that they did not succeed in infecting two dog-face
monkeys ( Cynocephalus species).
In the combined areas of Leopoldville, Boma, Matadi and the
Cataract Region, among a total of 1,172 persons examined, 8*8 per
cent, were infected, while of these latter 55 per cent, had been
diagnosed as cases of sleeping sickness. In the Gambia, the
previous Expedition had examined 1,043 natives, of which six only
harboured trypanosomes, and showed no definite symptoms.
Numerous lumbar punctures were made, and it was noted that
in many cases the trypanosomes never find their way into the
cerebro-spinal fluid, and in those cases in which they do they are
more likely to be found towards the termination of the disease; if
they gain access early in the disease, mania and other cerebral
symptoms are more likely to be prominent, but their entrance is in
no way correlated to the commencement of the fever or other
symptoms. In two later papers ( Memoir XVI, p. 97, and Memoir
XVIII, p. 1) on gland puncture in trypanosomiasis, the observers
favourably compared this with other methods of demonstrating the
presence of parasites. Following the work of Greig and Gray, they
concluded that by gland puncture, cases infected with trypanosomes
could be recognised at a much earlier period than hitherto. They
also, for the first time, observed a phenomenon frequently seen in
cases of trypanosomiasis, namely, auto-agglutination of the red
cells. The distribution of sleeping sickness in the Congo was
subsequently studied ( Memoir XVIII), it being concluded that the
increase during recent years was due, in a great measure, to the
increase in travel following the opening up of the country. In a
subsequent report ( Annals , Vol. I, p. 233), the trypanosomiasis of
cattle was dealt with. The investigators found that this disease
was very widely distributed in the Congo, the infecting parasite
being usually T. dimorphon. It was also observed that domestic
animals probably acquire a relative immunity to some strains of
trypanosomes, and may even recover spontaneously. Trypanosomes
were found in horses, mules and donkeys as well as in cattle, and
also in Tragelaphus scriptus .
Thomas, assisted by Linton and Breinl ( Memoirs XIII and
XIV), established by a long series of experiments that trypanosomes
found in (a) the cerebro-spinal fluid of Uganda sleeping sickness
cases, ( b ) the cerebro-spinal fluid and blood of Congo sleeping
sickness cases, (c) the blood of Congo ‘ trypanosome fever ’ cases, and
(d) the blood of Europeans infected in the Congo, were identical in
animal reactions and morphology with T. gambiense, Dutton. It
was also found (i) that the periodicity of the parasite is a prominent
feature, both in man and beast; (2) that the passing of a strain from
a susceptible into a very resistant animal does not attenuate the
organism, and that the morphological character is retained after
being passed through many hundreds of animals for nearly three
years; (3) that the parasites in an animal may sometimes become
more virulent, that such a strain may be particularly virulent for one
species of animal, and that the more rapid infection is not due to
the inoculation of a greater number of parasites than usual. In
addition to T. gambiense and T. dtmorphon, other pathogenic
trypanosomes were procured, and comparisons made between the
above organisms and T. evansi, T. brucei, T. equinum and
T. equiperdum. Cultivation of the different parasites was also
undertaken with success. Breinl gave a detailed account of the post¬
mortem changes in four cases of sleeping sickness ( Memoir XVI).
Extensive research was conducted into the treatment of trypano¬
somiasis, with the result that two drugs only were found to be of
any value in the disease, namely, arsenic and * Trypanroth'.
Thomas introduced atoxyl, a meta-arsenic anilin compound, a drug
which causes no pain on sub-cutaneous injection and may be
administered over a period of many months. He stated that it was
* the only remedy at present giving a prospect of a cure.’ Although
atoxyl would almost invariably cause the trypanosomes to disappear
from the peripheral blood yet since the parasites frequently
reappeared, it seemed possible that they might exist somewhere else
in the body of their host in a form uninfluenced by the' drug. Series
of experiments were, therefore, undertaken by Benjamin Moore and
Nierenstein, of the Bio-Chemical Department, and Todd. They
found that in the treatment of rats infected with T. brucei the
administration of atoxyl, followed by bi-chloride of mercury, gave
better results than treatment by atoxyl alone.
20
Moore, Nierenstein and Todd, continuing their researches on the
treatment of experimental trypanosomiasis ( Annals, Vol. II, p. 265),
found that in small animals, such as rats and rabbits, infected with
T. brucei , the results of treatment by atoxyl followed by mercury
salts were far superior to treatment by atoxyl alone; in large
animals, as donkeys, on the other hand, the combined treatment
was not found to be efficacious enough to be of practical value.
These workers also studied the effects of therapeutic agents on
trypanosomes in respect to (a) acquired resistance of the parasites to
the drug, and (b) changes in virulence of the strains after escape
from the drug ( ibid ., p. 221). Further bio-chemical research into
the subject was made by Nierenstein, who observed the acidity and
alkalinity of the blood in trypanosome infections, and found that
whereas the total acidity of the blood serum showed a marked
increase, the total alkalinity remained constant ( [ibid ., p. 227).
Later he made an extensive study of the chemo-therapeutics of
atoxyl (ibid., pp. 249, 323 and 329). Breinl conducted some
experiments on the combined atoxyl-mercury treatment of monkeys
infected with T. gambiense (ibid., p. 345), and demonstrated that
in five cases out of six, the administration of acetylated atoxyl and
sublimate, and Donovan’s solution, to monkeys (Cercopithecus
cdllithricus), effected a complete cure. As the result of Plimmer and
Thomson’s discovery of the trypanocidal action of antimony, Breinl
and Nierenstein investigated the action of aryl-stibinic acids in
experimental trypanosomasis (ibid., p. 365), and showed that both
p. and m. amino-phenyl-stibinic acids are fairly powerful trypano-
cides, the former being superior in its action. Owing to the
satisfactory results obtained in laboratory animals, a trial of the
former compound in sleeping sickness patients was advised, the dose
suggested being the same as for atoxyl.
In May, 1907, the Eighteenth Expedition of the School,
consisting of Dr. Allan Kinghom and Mr. R. E. Montgomery, was
despatched to Rhodesia and British Central Africa to study the
trypanosomiases of men and animals (Annals, Vol. II, pp. 53, 97,
333 and 387; Vol. Ill, pp. 259, 277 and 311). It was found that
sleeping sickness had already invaded N.E. Rhodesia, the first case
being seen in the Luapula division, adjoining the frontier of the
Congo. Glossina surveys were made, and the suggestion first
21
advanced that GL palpalis and Gl. fusca were not the only
transmitters of the disease, GL tnorsitans having also been observed
in infected areas. Gland palpations were made in 26,928 natives,
of whom 17*05 per cent, were found to have palpable glands. The
percentage of positive punctures was 77*7. The workers confirmed
the belief of Dutton and Todd that gland palpation, combined with
puncture, is a most useful measure, being a practical method of
isolating infected natives, and preventing any rapid extension of
the disease. It was found that there were between fifty and sixty
known cases of sleeping sickness in the country. The mode of
introduction and prophylaxis of the disease were studied, and
regulations drawn up by this Expedition were adopted and enforced
by the Government.
Concurrently with the research into human trypanosomiasis, the
Expedition pursued an enquiry into trypanosomiasis of domestic
stock in North-Western Rhodesia, in the course of which it was
established that the disease was very prevalent in that area, and
was due to T. dimorphon , T. vivax and a trypanosome morpho¬
logically allied to T. brucei . These trypanosomes could be
transmitted by GL tnorsitans , by Stomoxys calcitrans and by a
species of Lyperosia. The question of association of big game and
Gl. tnorsitans was considered, the opinion being that the distribu¬
tion of GL morsitans is entirely dependent upon the nature of the
country and its flora, that the association with the fauna is largely
fortuitous, and that a perpetual supply of mammalian blood is not
imperative, at least to its temporary existence.
The classification of trypanosomes was attempted, the observers
dividing the thirteen named species in the following way: —
T . theileri , T. equinum , T. gatnbiense and T. equiperdum , easily
recognised by their morphology or animal reactions; the nine
remaining species sub-divided into three groups having as
their types:—(1) T . evanst (with T. brucei and T . sudanense ).
(2) T . dintorphon (with T . congolense and T . pecaudi , and
(3) T . nanunt (with T. vivax and T. cazalboui ).
Research was undertaken into many other points, and included
observations on the parasites occurring in the intestine of GL palpalis
and in the intestine and proboscis of Gl . tnorsitans .
The many problems of trypanosomiasis continued to occupy the
22
energies of the staff at Runcorn. Yorke took up the study of
immunity in trypanosome diseases. He was able to prove ( Annals,
Vol. Ill, p. 565) that a diminution of the haemolytic complement
only takes place in the last stages of the disease, and that, in an
experimental case, the complement had practically vanished shortly
before the death of the animal, when the blood was swarming with
parasites. He also worked on the protective action of the serum of
animals in a state of chronic infection or immune to various kinds
of trypanosomes. Later, he investigated the condition of the blood
which gives rise to the phenomenon of auto-agglutination of the red
blood cells in animals infected with trypanosomes and to sleeping
sickness in man ( Proc. Roy. Soc., Ser. B, Vol. LXXXIII, p. 238,
and Annals , Vol. IV, p. 529). Experiments showed that auto-, iso-
and hetero-agglutinin exist in the blood of many normal animals,
and are frequently present in much greater amount In the blood of
infected animals. Apart from infection with trypanosomes, well
marked auto-agglutination was found to be an extremely rare
phenomenon. Ross and Thomson studied a case of sleeping
sickness and demonstrated a regular periodical increase of the
parasites {Annals, Vol. IV, pp. 261 and 395). These workers
confirmed the claim that there is a life cycle of trypanosomes in the
vertebrate host {ibid., p. 465). In 1910, Stephens observed a
marked peculiarity in the morphology of a trypanosome from a rat
supposed to be infected with T. gambiense. This parasite was
described by himself and Fantham {Annals, Vol. IV, p. 343), the
animal reactions observed by Yorke {ibid., pp. 351 and 385), and a
new species was founded, to which the name T. rhodesiense
Stephens and Fantham was given, based on the peculiar posterior
position of the macronucleus. It was later established that T. brucei
exhibits this peculiarity.
In 1911, the Twenty-seventh Expedition of the School, consisting
of Dr. J. L. Todd and Dr. S. B. Wolbach, was despatched to the
Gambia to investigate sleeping sickness. Reports of the work done
were published in the Annals, Vol. V, p. 245.
In the course of this investigation, 12,298 persons were palpated,
and the observers put on record their opinion that gland palpation
and puncture was by far the best procedure for the diagnosis of
trypanosomiasis. It was found that at least o’8 per cent, of the
Annals Trap, Med Parasiml.. Vol. XV
PLATE 1
C. Ti filing & Co. , Lid.. Imp .
*3
population of the Gambia was infected with trypanosomes. Strong
recommendations were made for the control of the disease in the
Gambia, including a continued examination of the whole population,
the establishment of villages for isolation, observation and treat¬
ment of cases, and the appointment of a special staff for the
administration and execution of these projects. Stannus and
Yorke examined in rats the parasite from a case of sleeping sickness
contracted in Nyasaland (ibid,, p. 443), and were convinced that
the trypanosome in question was not T. gambiense but probably
identical with T. rhodesiense , which it resembled very closely,
having a posterior nuclear form. A study was made by Yorke and
Blacklock of the trypanosomes from a horse naturally infected in the
Gambia (ibid., p. 413). The parasites consisted of a long form
with a free flagellum, and also a short form without a free flagellum.
The former was considered, from its morphological appearance and
animal reactions, to belong to the T. vivax group; the latter form
was subsequently identified (Annals, Vol. VI, p. 107) as
T. dimorphon, sensu Laver an and Mesnil. Blacklock measured
one thousand examples of this form, and found that the average
length was 13*3/* (Annals, Vol. VI, p. 287). Measurements of one
thousand examples of T. vivax in goats were made by Blacklock,
the average length being 27*7/1 (Annals, Vol. V, p. 521). He also,
using the same strain of T. vivax, measured fifty trypanosomes
drawn from a rat and a rabbit respectively. In the former the
average length was 21 *i/*, in the latter 20*8/1 (ibid., p. 537).
In 1911, at the request of the British South African Co.,
Dr. Kinghom and Dr. Yorke were sent to Rhodesia (Luangwa
Valley) to study sleeping sickness. It was quickly placed beyond
doubt that Gl. morsitans was the carrier of the human trypanosome
(Annals, Vol. VI, p. 1). The investigators inoculated rats from
twelve cases of human trypanosomiasis, eleven of which occurred in
villages in the Luangwa valley. In every instance they observed
the posterior displacement of the macronucleus, characteristic of the
trypanosome described by Stephens and Fantham; the animal
reactions agreed in all respects with those obtained from infection
with T. rhodesiense. Elaborate transmission experiments with
both wild and laboratory-bred Glossina morsitans were successfully
carried out in rats and monkeys; the duration of the cycle in the
H
fly (approximately fourteen days) was found to be shorter than in
experiments of previous investigators with Gl. palpalis and
Gl. morsitans. It was observed that an infected fly retains the
power of transmitting the disease during its life, and is infective
at each meal, but that mechanical transmission does not occur if a
period of twenty-four hours has elapsed since the infecting meal.
Certain species of buck, viz., waterbuck, hartebeest, mpala and
warthog, were found to be infected with the human trypanosome,
as well as a native dog. Later {Annals, Vol. VI, p. 269), it was
found that 16 per cent, of the local game were infected with
T. rhodesiense, and it was established that the game and fly
strains were identical with the human trypanosome. In all, six
species of tryanosomes were found in game and domestic stock
in the Luangwa Valley {Annals, Vol. VI, p. 301), namely,
T. rhodesiense and T. pec or urn, transmitted by Gl. morsitans and
probably by insects other than tsetse-flies; T. vivax and
T. nanutn, probably transmitted by Gl. morsitans; and two others,
one of which was possibly T. montgomeri. At least 37’5 per cent,
of the buck were found to harbour pathogenic trypanosomes. In a
later report {ibid, p. 317) a new trypanosome, T. ignotum,
infective to monkeys and a rabbit, was described. The vertebrate
host was not discovered. This trypanosome is now known as
T. simiae, having been previously found in the same year by the
Royal Society S.S. Commission. Still later {Annals, Vol. VII,
p. 254), descriptions were given of T. multiforme, sp. n., and of
T. tragelapki, the latter closely resembling T. in gens. In the course
of experiments in the development of T. rhodesiense in Gl. morsitans
{Annals, Vol. VI, p. 405), it was observed that the cycle was
influenced to a marked degree by the temperature to which the flies
were subjected, high temperatures (75 0 to 85° F.) being favourable
and low ones (6o° to 70° F.) unfavourable to the development of the
parasites. Parasites might persist in the fly at at incomplete stage
of their development for at least sixty days under unfavourable
climatic conditions. It was found {Annals, Vol. VI, p. 495) that
in every fly capable of infecting animals with T. rhodesiense the
salivary glands were invaded, also that on every occasion on which
the salivary glands were infective the trypanosomes in the intestines
were virulent. Invasion of the salivary glands was only observed
*5
in the case of flies infected with T. rhodesiense and not in the case
of any other trypanosomes met with in the Luangwa Valley
or on the Congo-Zambesi watershed. It was calculated that
approximately 3‘5 per cent, of the flies might become permanently
infected and capable of transmitting the virus {Annals, Vol. VII,
p. 183). The chief reservoir of the human trypanosome was found
to be the antelope. Stephens and Fantham {Proc. Roy. Soc., B,
Vol. LXXXV, p. 223, and Annals, Vol. VI, p. 131) made a
bionomic study of T. rhodesiense. One thousand specimens were
measured, and it was found that the average length was 2 y 6 /t, as
compared with T. gambiense 22'in, and T. brucei 2 y 2 /i. The
average length of T. rhodesiense in man and other species of animals
was ascertained. Those in the rat were found to be the longest
(24*5 n) and those in the rabbit the shortest (ig‘4/»). Of the three
species T. rhodesiense was found to be richest in long and poorest in
intermediate forms. Another series of measurements was under¬
taken {Annals, Vol. VII, p. 27), when each day, for the first ten
days of infection, one hundred trypanosomes from the same rat
were measured, the results being again compared with T. gambiense.
It was found that the day of infection was of great importance, as
there was a great variation in the percentage of ‘ stumpy * forms on
different days, e.g., 53 per cent, on the seventh day to 5 per cent,
oh the tenth day. The larger the sample of trypanosomes taken,
the smaller the variation in the average length.
J. G. Thomson succeeded in cultivating T. rhodesiense by the
use of a modification of the Novy-MacNeal-Nicolle medium. The
changes taking place in the.trypanosomes were described {Annals,
Vol. VI, p. 103), and it was noted that when development was
rapid two distinct types could be distinguished on the fourth day.
In cultures which develope more slowly the trypanosomes disappeared
about the third or fourth day, reappearing about the sixth, and on
the eighth day spirillar forms were seen to be splitting off.
Differentiation into the so-called ‘ male ’ and ‘ female ’ forms took
place during the eighth, ninth and tenth days. Inoculation of
animals from the cultures was unsuccessful. In a subsequent
investigation with J. A. Sinton {ibid., p. 331), Thomson
successfully cultivated T. gambiense as well as T. rhodesiense, the
former for a period of thirty-seven days, the latter for twenty-one
26
days only. The life history of these trypanosomes in culture tubes
was similar to that occurring in the gut of the insect host. The
cultures lost their infectivity after the third day, and it was
suggested that probably their transference to a new medium or
environment similar to that of the salivary glands of the tsetse fly
might be required to permit the full history of the trypanosomes
being completed. Blacklock observed the vitality and changes
undergone by trypanosomes in the cadaver of the animal host
{ibid., p. 55). He found that T. gatnbiense and T. rhodesifnse
can remain infective in the blood of the dead animal for forty-eight
hours. Blacklock also made a study of the posterior nuclear forms
of T. rhodesiense in rats {Annals, Vol. VII, p. 101). He found
that they first appeared in the blood from the sixth to the tenth
day of the disease, in a count of a thousand trypanosomes, that
they increased in numbers in the later stage of the disease, and
that they increased relatively to other forms of trypanosomes. They
showed definite powers of resistance to disintegration in the cadaver
of the animal host. It was suggested that such forms might occur
as a constant constituent of certain strains.
Stephens and Blacklock made a morphological study of
T. brucei (the Zululand strain) and of the trypanosome of the same
name from the Uganda ox {Annals, Vol. VII, p. 303). They
asserted the non-identity of the two strains, and proposed for the
latter the name T. ugandae. Blacklock and Yorke {ibid., p. 603),
studying the pathogenicity of T. congolense (Broden) and T. nanurn
(Laveran), came to the conclusion that they were the same parasite.
Yorke and Blacklock studied the characters of the more
important mammalian trypanosomes {Annals, Vol. VIII, p. 1), and
compiled a convenient table of the main differential points. The
value of the cycle of the trypanosomes in their invertebrate hosts as
an aid to differentiation of species was also put forward.
Todd re-examined the flagellate found by Dutton in 1902 in
the blood of Gambia house mice, and came to the conclusion that
it was T. acomys {ibid., p. 469).
Yorke and Blacklock experimented with antimony trioxide in
the treatment of experimental trypanosomiasis {ibid., p. 55),
controlling and extending some of the work of Kolle and others on
the use of this drug. Various strains were used in the experiments.
It was found that small animals could withstand a relatively much
larger quantity of the drug than large ones. Post-mortem evidence
proved that the proportion absorbed during a period of six months
was exceedingly small. A certain number of cures would seem to
have resulted, as several animals remained negative without relapse
for over two hundred days, and sub-inoculated animals were not
infected. Most strains appeared very susceptible to the drug, but
X. gantbiense and X. lewisi proved refractory.
Seidelin tested the effect of salvarsan-copper on white rats
infected with a strain of trypanosomes of the X. brucei group kept
in guinea-pigs and rats in West Africa (Annals , Vol. IX, p. 197).
The best results were obtained with the injection of a dose of
0*0064 gm.; in such a case the trypanosomes disappeared from the
blood on the following day and remained absent for fifteen days,
death occurring on the twenty-eighth day; in several other cases
the life of the animals was prolonged for a few days more.
The Thirty-second Expedition of the School, consisting of
Drs. Yorke and Blacklock, was despatched to Sierra Leone in 1914.
Research was undertaken into the bionomics of G. palpalis in Sierra
Leone, with special reference to its pupal habits (Annals, Vol. IX,
p. 249). It was found that the breeding-grounds of G. palpalis
are not so strictly limited to the immediate vicinity of water as had
hitherto been believed. Mangrove swamps do not constitute a
breeding-ground. The pupae do not hatch when subjected to daily
flotation cn sea water. The ground around the trunk of oil palms
which have not been stripped of their lower petioles constitutes an
excellent breeding-place for G. palpalis ; they can breed in localities
in which practically the only tree is the oil palm. A study of the
food of G. palpalis in the Cape Lighthouse Peninsula, Sierra Leone
(ibid p. 363), showed that about 8 per cent, of the wild G . palpalis
in that district contained recognisable red blood cells—7 per cent,
of mammalian origin and 1 per cent, nucleated red cells of unknown
origin. Neither shed blood nor other fluid which is exposed (not
covered by a membrane) can be imbibed by G. palpalis . Fluids
such as solutions of sugar, sodium chloride, etc., protected by a
membrane (e.g., thin rubber sheeting), were taken up, but less*
quickly and readily than blood. It was thought that in nature
G. palpalis may, under certain conditions, take up fluids other than
28
blood. The human trypanosome ( T . gambiense) was discovered in
an ox in Sierra Leone (ibid., p. 383), thus demonstrating that
domestic stock forms a reservoir for the virus of sleeping sickness.
Among other animal parasites found in domestic stock in Sierra
Leone (ibid., p. 413), T. congolense and T. vivax were most
commonly encountered. Trypanosomiasis of cattle was common.
Of one hundred and forty-three examined, trypanosomes were found
in nineteen after a single examination.
In 1919, Escomel recorded the discovery of trypanosomes in the
blood of a patient coming from Peru, which he described and
considered to be Schizotrypanum cruzi. YorEe, examining this
description, was led to doubt, on morphological grounds, whether
the identification was correct, and proposed for this Peruvian
parasite the name T . escomeli , in recognition of its discoverer. A
feature of the symptomatology of the case was the overpowering
somnolence from which the patient suffered, a symptom not hitherto
noted in Chagas’ disease, although a striking feature of the African
trypanosomiasis of man (Annals, Vol. XIII, p. 459).
Macfie described a trypanosome found in the blood of a snake,
Naja nigricollis , in the Gold Coast, resembling in some degree
T. primeti, but differing from it in size and proportion. He
proposed for it the name T. voltariae (Annals, Vol. XIII, p. 23).
YELLOW FEVER
In 1900, the Fourth Expedition, consisting of Drs. H. E.
Durham and Walter Myers, went to Brazil to study yellow fever.
Dr. Myers died of the disease in January, 1901, while Dr. Durham,
who also contracted it, recovered and published a report of the work
of the Expedition (Memoir VII) in 1902.
The U.S. Yellow Fever Commission in Cuba had not yet
established the transmission by Stegomyia when the Liverpool
Expedition set to work at Para; for which reason the course of their
investigation was chiefly directed at first to the search for
some protozoal parasite. Only seventeen autopsies on yellow fever
cadavers were obtainable. An extremely slender filiform bacillus
was observed to occur constantly in the tissues and intestines, and
much time was spent in isolating it and in attempts at cultivation.
29
Dissection of specimens of Culex fatigans captured in suspected
houses showed large numbers of a similar bacillus. Owing to the
death of Dr. Myers, the research came to an end.
In 1905, Professor Rubert Boyce was despatched by the School
to New Orleans, to observe the work of the U.S. Medical Authorities
in dealing with the outbreak of yellow fever there. In Memoir XIX
Boyce gave an account of the vigorous campaign which was
successful in ridding the city of Stegomyia. Subsequently, at the
request of the Colonial Office he visited Honduras to make a
report on the conditions existing in that Colony with reference to a
recent outbreak of yellow fever (Waterlow & Sons, 1906).
In April, 1905, a second Yellow Fever Expedition was sent to
the Amazon, the members being Dr. Wolferstan Thomas and
Dr. Breinl. A permanent laboratory was established so that
patients could be kept under continued observation, and experiments
inaugurated which would have been impossible under other
conditions. Series of reports by Thomas were published in
the Annals, Vol. IV, p. 1. It was recorded that the most serious
disease to which the foreign population was liable was yellow fever,
and this disease was made the object of extensive investigation.
‘ Chimpanzees have been successfully inoculated; rabbits and guinea-
pigs exhibited certain reactions when inoculated with infective
blood from yellow fever cases or subjected to the bites of infected
S. calofus.' (See also Trans. Soc. Trop. Med. and Hyg., Vol. Ill,
p. 59.) The resources of the laboratory in Manios were placed at
the disposal of the State, the doctors, the hospital and the poor of
the city. Examinations of blood, agglutination reactions for
typhoid and paratyphoid, bacteriological examinations of water
and milk, post-mortems and pathological reports were made, and
the laboratory so conducted as to be of the greatest possible service
to the community. Papers on ‘Oesophagostomiasis in Man’ and
the condition known as ‘ Mossy Foot’ were published in the Annals,
as also an article on the ‘ Mosquitoes of the Amazon region ’ by
Newstead jointly with Thomas.
In 1910, at the request of the Colonial Office, Professor Boyce
was despatched to the Gold Coast and Sierra Leone, to report on
the outbreak of yellow fever at those places (Trans. Soc. Trop. Med.
and Hyg., Vol. IV, p. 33). It was observed that the so-called
3 °
classical type of yellow fever was comparatively rare amongst native
races, and the reason was advanced that natives are partially
immunised by being bom and brought up in an endemic area
(Annals , Vol. V, p. 103). Those removed in childhood from such
an area become non-immune, and therefore liable to succumb to an
epidemic.
In 1911, the study of yellow fever was taken up with energy by
Professor Boyce, who suggested and superintended the establish¬
ment of a Yellow Fever Bureau at the Liverpool laboratories. A
Bulletin was issued, in which the Director of the Bureau, Dr. Harald
Seidelin, and other investigators published the results of their
researches. Abstracts of reports from all over the world were also
made, and reviews published of current literature on yellow fever
and allied subjects, including pappataci fever and dengue.
Research was undertaken into the etiology, diagnosis and treatment
of yellow fever. Seidelin observed the occurrence of protozoon-likc
bodies in the blood and organs of yellow fever patients ( t fourn . Path .
and Bad ., Vol. XV, p. 282). At a meeting of the Society of
Tropical Medicine and Hygiene, in January, 1911, these organisms
were demonstrated in preparations of the blood and sections of the
kidney, and later ( Yellow Fever Bulletin , Vol. I, p. 229), Seidelin
described their morphology, and proposed for them the name of
Paraplasma -flavigenum , g. n., sp. n. In December, 1911, he was
despatched to Yucatan, where an epidemic of yellow fever was
raging; a report of this Expedition was published in Yellow
Fever Bull ., Vol. II., p. 123. THe total number of cases officially
diagnosed during the outbreak was seventy-three, with thirty-eight
deaths. The existence of a ‘microbe carrier’ was put forward as
being responsible for the maintenance of the virus during periods
when the disease is latent, it being suggested that the fragile
infected Stegomyia could not be the lasting reservoir. It was also
considered that one attack does not necessarily confer immunity.
The usually mild form of the disease in natives was said to be due
either to an unrecognised infection in childhood or to the hereditary
transmission of anti-bodies from immune parents. Clinical features
of the cases observed were given. Paraplasma -flavigenum was seen
in fifteen confirmed cases of yellow fever out of sixteen examined.
Summing up the results of his blood examinations for P. flavigenum
3 *
in undoubted cases of yellow fever, Seidelin found that on this and
former occasions he had approximately one hundred and six
positive cases out of a total of one hundred and twenty. Records
were made of the examination of four hundred and twenty-one
specimens as controls. In one hundred and thirty-six cases malarial
parasites were found, and in two hundred and eighty-three no
parasites at all, while in two young children under no suspicion of
yellow fever P. flavigenunt occurred. It was suggested that these,
together with two others previously observed, might be 4 microbe
carriers.* Blood from yellow fever cases was inoculated into four
guinea-pigs, but none showed symptoms resembling the disease.
A mosquito survey of Merida was made, and recommendations
advanced as to methods of extermination. A section of Seidelin*s
report was devoted to a reply to those who had criticised his work.
In 1915, the Yellow Fever Commission (West Africa) concluded
that no proof had been given that P . flavigenum was of a protozoal
nature, and that the nature of the virus of yellow fever still remained
undetermined.
In December, 1912, Seidelin was sent to Jamaica in order to
investigate the nature of the disease called 1 vomiting sickness,*
prevalent in that island during the winter months, and responsible
for a considerable mortality, chiefly among native children. A
report of this Expedition was published in the Annals (Vol. VII,
p. 377) and also in the Yellow Fever Bulletin (Vol. Ill, p. 7).
Sixty-two cases were observed, but no causal organism was recog¬
nised ; it was concluded that ‘ vomiting sickness* was a local disease
and could not be accepted as a form of meningitis, which view had
been advanced by Scott two years before the latter*s demonstration
of its true origin.
RELAPSING FEVER AND SPIROCHAETES
By the death of Dr. Dutton from tick fever while serving with
the Congo Expedition, the School suffered the loss of one of its most
brilliant workers, who, although only twenty-nine years of age, had
already won a recognised position throughout the scientific world.
Towards the end of 1904, Dr. Dutton and Dr. Todd had reached
Stanley Falls, and they were able to demonstrate independently the
32
cause of tick fever in man, a discovery made a few weeks previously
by Ross and Milne in Uganda. Further, they were able to prove
the transference of the disease from man to monkeys by means of a
particular species of tick. In Todd’s report of this discovery
(Memoir XVIII) clinical descriptions were given of twelve native and
two European cases (the last two being those of the investigators
themselves), all of whom were infected with a spirochaete thought
at first to be 5 . obermeieri. A number of inoculation experiments
with laboratory animals were carried out, it being found that to
monkeys • alone the parasite appeared to be uniformly pathogenic.
Observations were made upon the distribution and bionomics of the
human tick in the Congo, and included in the report was a
description of Ornithodoros moubaia by Professor Newstead.
Research was proceeding at Runcorn upon material brought back
from the Congo by Dr. Todd. Dr. Breinl, invalided home from the
Fifteenth Expedition, carried out with Dr. A. Kinghom extensive
studies on ‘tick fever’ and ‘relapsing fever.’ Observations were
made on the animal reactions of the spirochaete discovered in the
Congo cases of human tick fever, and brought to England in infected
monkeys and ticks. In the course of these experiments, infection
was produced not only in monkeys, but also in a horse, a dog,
rabbits, guinea-pigs, rats and mice. This fact caused the observers
to conclude that the organism was distinct from Spirochaeta
obermeieri , pathogenic hitherto to monkeys only. Further experi¬
ments were undertaken, confirming this conclusion; and the new
species was given the name of Spirochaeta duttoni (Breinl and
Kinghom, 1906, Memoir XX). Studies were made of this organism
(Memoir XXI), in the course of which a clinical comparison was
made between African tick fever and European relapsing fever, and
the research into the animal reactions of this spirochaete in various
animals amplified and completed. Experiments in immunity were
continued, the conclusions reached being : (1) In animals which have
recovered from the infection there is a relatively active immunity
of comparatively long duration; (2) immune serum cannot produce
passive immunity, nor has it any curative action; (3) hyper-immune
serum does not protect a susceptible animal, nor does it prevent
relapse, but it mitigates the severity of the infection and occasionally
cuts short an attack; (4) there is a slight inborn immunity of short
33
duration. It was also shown that 5 . duttoni may pass in utero
from mother to foetus, and extensive studies were carried out to
determine the role played by the spleen in infection by spirochaetes.
Attempts were made to transmit spirochaetes by the bites of Cimex
lectularius , but without success. Subsequently ( Annals, Vol. I,
p. 435) Breinl studied the morphology and life history of 5 . duttoni ,
while Markham Carter (ibid ., p. 15) described the multiplication
and important changes in form of S. duttoni in eggs laid by infected
ticks. Still later (Annals, Vol. V, p. 479), Fantham studied the life
cycle of spirochaetes, amongst those considered being 5 . duttoni ,
S. recurrentis and S. marchouxi . Subsequently (Annals, Vol. VIII,
p. 471) he investigated the granule phase of the parasite, a detailed
study being given, while serving with the Expedition to Khartoum,
to 5. bronchialis , in which it was found that the granules formed
by the spirochaete were the cross infective stages of the organism
(Annals, Vol. IX, p. 391).
In 1917, while making microscopical examinations of stained
smears from the stools of five hundred and fifty-four patients,
admitted to hospital for dysentery, Carter found that 56*5 per cent,
were infected with Spirochaeta eurygyrata . A control investigation
on one hundred cases free from intestinal disorders showed 41 per
cent, to be infected (Annals, Vol. X, p. 391). Repeating this
investigation amongst a normal population, Macfie and Carter
(Annals, Vol. XI, p. 75) examined eighty-two hospital patients
suffering from some surgical condition, and twenty-three normal
healthy men. None of the cases had ever resided in the tropics.
Of the hospital patients 56*2 per cent., and of the healthy men
43*8 per cent., harboured 5 . eurygyrata . A second species of
spirochaete was discovered in the intestine of one case, which, owing
to its larger size and certain morphological peculiarities, was
considered to be a new species, and named by them Spirochaeta
intestinalis . Macfie and Yorke examined the morphology of the
spirochaetes responsible for European, African and Indian relapsing
fevers (Annals, Vol. XI, p. 81), and reached the conclusion
that there is at present no means of distinguishing these parasites
morphologically.
34
AMOEBIASIS
Research into the amoebae parasitic in the human intestine was
undertaken by Fantham, and a study commenced of the life history
of E . coli as seen in cultures {Annals , Vol. V, p. ill).
Carter, Mackinnon, Matthews and Smith conducted extensive
researches into the protozoal findings in cases of amoebic dysentery.
In their first report {Annals , Vol. X, p. 411) they recorded the
results of four thousand three hundred and thirty-four examinations
of nine hundred and ten patients suffering from this condition.
Protozoal infections were discovered in 44*2 per cent.; E. histolytica
was found in 10*3 per cent, of the cases; E. coli in 25*4 per cent.;
G. intestinalis in 18*6 per cent.; T. intestinalis in V2 per cent.; and
C. mesnili in 2*7 per cent. Their second report {Annals , Vol. XI,
p. 27) recorded similar examinations of one thousand seven hundred
and thirteen cases of dysentery. Stress was laid upon the necessity
for repeated examinations of each patient, as cases found negative
the first and second times may prove on further examination to be
E . histolytica carriers. The subject of ‘negative periods* (absence
of vegetative forms and cysts) in infected cases was also dealt with.
A third report of this investigation appeared in Annals , Vol. XIII,
p. 83. Yorke and the above-mentioned observers examined for
intestinal protozoa three hundred and forty-four persons who had
never been out of England ( Annals , Vol. XI, p. 87). Of this
number, two hundred and six were healthy young men of about
18 years of age who had recently entered the Army. A single
examination of each of these cases revealed the interesting fact that
3*9 per cent, were infected with E. histolytica . In an address to
the British Medical Association {B.M.J. t April 12th, 1919), Yorke
emphasized the importance of discovering whether the infection in
this country is recent or otherwise. He was inclined to believe that
it was not recent because (1) carriers must have frequently entered
this country before the war; (2) all the necessary factors for the
spread of the infection are to be found in this country; (3) there are
authentic records of cases of amoebic dysentery and liver abscess
before 1914.
Stephens and Mackinnon treated eighty-one cases infected with
E. histolytica with ‘ alcresta ipecac,’ an adsorption compound of
.7 rrrtals Trap. Med. & ParasitolVol. XI*
PLATE Fill
MAIN LABORATORY
C. Tittliio <o? Cn„ Ltd,, lw[>.
35
emetine and aluminium silicate ( Annals , Vol. X, p. 397), with the
result that about two-thirds of the patients were freed from amoebic
cysts. Carter and Matthews, using Cropper and Row’s method of
concentration (Annals, Vol. XI, p. 195), found E. histolytica cysts
in five of one hundred and thirty-three apparently negative cases
which had already received three ordinary routine microscopical
examinations. Smith made a mensurative study of the cysts of
E . histolytica and E. coli (Annals , Vol. XII, p. 27), and investi¬
gated the question of the number of races in the former parasite
(Annals, Vol. XIII, p. 1). It was shown that not all infections of
E. histolytica remain constant from one day to another in the
average size of their cysts. The species can be divided into two
races characterised by larger and smaller cysts, respectively. Infec¬
tions with E . histolytica in healthy carriers who have never been out
of England were shown to be characterised by a smaller proportion
of the ‘ small ' race, and also by a reduced proportion of the larger
cysts of the ‘ ordinary * race, as compared with infections from
convalescent dysenteries from abroad. Investigating the incidence
of amoebic dysentery in asylum patieiits never out of England
(Annals, Vol. XIII, p. 177), Smith found that of five hundred and
four patients examined, fifty-nine had acute dysentery, and in three
cases vegetative E . histolytica were found in the stools.
BER1-BERI
Simpson and Edie undertook research into the relation of the
organic phosphorus content of various diets to diseases of nutrition,
particularly beri-beri. After a review of the work of Schaumann
and others (Annals, Vol. V, p. 313), the investigators recorded their
own experiments with pigeons fed on various kinds of rice, white
bread and whole-meal bread. A study was made of the anti-
neuritic bases of vegetable origin in relationship to beri-beri, the
properties of the yeast extracts investigated, and a method adopted
for the isolation of torulin, the anti-neuritic base of yeast (Annals,
Vol. VI, p. 235).
36
HELMINTHIASIS
Stephens ( Thompson-Yates Lab . Reports , Vol. VII, p. 9),
described the morphology of Gastrodiscus hominis , of two new
human cestodes, Dibothriocephalus parvus and Taenia brentneri,
of a new linguatulid, Porocephalus pattoni {Annals , Vol. I, p. 549 )*
and of a new human nematode, Strongylus gibsoni {Annals, Vol. II,
p. 315). Observations on the hooklets of Cysticercus cellulosae in
man {Annals, Vol. II, p. 391) showed that there is an irregularity
of development affecting both the number of the hooklets and, more
especially, the size. A fluke, found by Newstead in the alimentary
canal of a Nicaraguan turtle, was described by Stephens {Annals,
Vol. V, p. 497), who proposed for it a new genus and named it
Desmogonius desmogonius . The fluke from the liver of native dogs
at Kasauli, India, was separated by Stephens from the genus
Opisthorchis owing to the existence of a process or pedicle bearing
on its summit the genital opening and a ventral sucker, and placed
in a new genus as Par opisthorchis caninus {Annals, Vol. VI, p. 117).
Breinl and Hindle described a new Porocephalus, found in the lung
of one of their experimental monkeys, and distinguished from the
known species by the presence of an appendage on the outer pair
of hooks only. They proposed for it the name of Porocephalus
cercopitheci {Annals, Vol. II, p. 321). Dogs in Freetown were
found by Yorke and Blacklock to be heavily infected with Ankylos-
toma caninum and A . ceylanicum, the species being present in about
equal numbers {Annals, Vol. IX, p. 425).
In 1916, Stephens contributed the section on Helminths to
‘ The Animal Parasites of Man,’ issued jointly with Fantham and
Theobald.
In 1917, at a Veterinary Hospital attached to a remount depot
in the neighbourhood of Liverpool, Yorke and Macfle started an
investigation into the parasitic worms causing a heavy mortality
amongst hcrses recently imported from America. The parasites
belonged for the most part to various genera of the family
Strongylidae, and in the course of their study Yorke and Macfle
described eight new species and one new variety, viz., Cylicostomum
longibursatum , C . minutum, C. pseudocatinatum, C . pater at um ,
C. tridentatum , C. triramosum , Cylindro pharynx rhodesiense,
37
Gyalocephalus equi and Cyhcostomum nassatum, Looss, var.
parvum (Annals, Vols. XI-XIV).
Yorke and Southwell described a nematode from the intestine
of a zebra, which certain minute characters of the head, and
also the position of the vulva, led them to regard as a new
species, for which they proposed the name Crossocephalus zebrae
{Annals, Vol. XIV, p. 127).
FI LARI AS IS
The second part {Memoir IV) of the Report of the Third
Malarial Expedition was devoted almost entirely to Filariasis.
Eight new species, found during the examination of a large number
of West Afrcan birds, were described, namely:— F. cypseli,
F. spiralis avium, F. fusiformis avium , F. spiralis avium major,
F . falciformis , F. bibulbosa, F . capsulata and F. shekletoni .
Observations were also made on human filariasis in West Africa, it
being found that throughout the whole of that area the natives
appeared to be infected with F . nocturna, diurna and perstans.
With regard to the two first species, the majority of the cases
encountered were atypical, in that, embryos were either never absent
from the peripheral blood, or the maximum did not occur at mid¬
day and mid-night or thereabouts according to the species. Among
the former cases there were many showing decided periodicity, and,
among the latter, the hour at which the maximum number was
present varied considerably. In some cases, two maxima during
the twenty-four hours were indicated. In the examination (day
blood) of three hundred and ninety natives of all ages up to about
eighteen years, one case only, aged eleven years, was infected. The
observers succeeded in infecting A . costalis (proboscis) with
F . nocturna . They considered that the weight of evidence was on
the side of the identity of F . nocturna and F . diurna , but that many
points remained to be cleared up before the question could be
settled.
A study of the periodicity of Microfilaria nocturna was made by
Yorke and Blacklock {Annals, Vol. XI, p. 127). They found that
obstruction to the passage of M. bancrofti through the cutaneous
vessels occurs at all times of the day and night, but is at a minimum
3 »
at the end of the period of bodily activity. Although this obstruc¬
tion aids in the piling up of the larvae in the cutaneous vessels, it
is in no way responsible for the nocturnal periodicity, which is
primarily dependent upon periodic variations in the arterial supply
of larvae to the cutaneous vessels. By reversing the hours of sleep
and activity, cutaneous immigration gradually becomes diurnal
instead of nocturnal, the complete inversion of the periodicity being
accomplished in from four to eleven days. The number of
microfilariae, as judged from the maximum concentration in the
cutaneous blood, remained at practically a constant level during the
period of observation. The number of microfilariae in the urine
varied greatly, the variation giving no indication of either a
nocturnal or a diurnal periodicity. The number of microfilariae in
the renal and vesical' vessels exhibited a nocturnal periodicity
analagous to that in the cutaneous vessels.
ENTOMOLOGY
In 1907, Professor Newstead, jointly with Drs. Dutton and
Todd, issued a report (Annals, Vol. I, p. 1) upon the insects and
other arthropoda collected in the Congo Free State. Among the
new genera and species described and figured were Eretmapodites
inornatus, Stegomyia luteocephala , S. albomarginata , Duttonia
tarsalis , D. africana , Culex laurenti , C. par , Mimotnyia africana ,
M . malfeyti, Boycia mimomyiajormis , Haematopota duttoni ,
H . trimaculata , Tabanus billing! oni> Glossina maculata and
Stomoxys omega • The habits and structural characters of the
larva of Simulium ornaturn (a European species) were described.
They occur on the under sides of submerged leaves or blades of
grass in those parts of a stream which are rapidly flowing and fully
exposed to the sun. Their mode of progression resembles that of
looper caterpillars, spinning a network of silken threads along
which they travel. The period of pupation varies from two to six
days. The imagines escape through a slit in the thorax, and
occasionally may be seen completely immersed in water with their
wings folded so as to encase an air bubble. The distribution of
tsetse-flies in the Congo was recorded and illustrated by a map,
together with observations on the bionomics of the flies. The life
history of Stomoxys calcitrans was also described in detail.
Annals 7 rop. M(d if Parasito!., Vol. XV
PLATE IX
ENTOMOLOGICAL DEPARTMENT
C. Titiling if Co., Ltd., Imp.
39
In the following year, Newstead made a study of the bionomics
of the common house-fly (- Annals, Vol. I, p. 507). The chief
breeding-places were determined, and it was established that the
life-cycle of the fly, in all kinds of fermenting material, is reduced
to the minimum period of ten to fourteen days; and that in the
absence of such artificial heat the cycle may occupy a period from
three to five weeks or more, according to the temperature of the
outside air. It was found that house-flies do not depend entirely
upon excessively warm weather for breeding purposes. Methods of
prevention were suggested, and some notes appended on other
insects found during the investigation.
In 1908, Newstead went to Jamaica to study cattle ticks. In
the course of the investigation, twenty-five estates were visited and
the cattle inspected in each, and large numbers of ticks were
collected in every district {Annals, Vol. Ill, p. 421). Methods of
treating tick-infected stock were made the subject of enquiry, and
whenever possible practical demonstrations were given. The nature
and extent of injury caused by insects and other pests to cultivated
crops was studied, and methods of control advised. Descriptions
were given of the nine species of ticks found in Jamaica, namely,
Argas persicus, Margaropus annulatus australis , Rhipicephalus
bursa , R. sanguineus , Dermacentor nitens , Amblyomnta maculatum ,
A . cajanense, A. dissimile, Aponomma sp.
In 1906, Newstead investigated the life history of Stomoxys
calcitrans (Journ. Econ. Biol., Vol. I, p. 157) and, together with
Stephens, described the anatomy of the proboscis of Glossina
palpalis {Memoir XVIII, p. 53), and subsequently ( Annals , Vol. I,
p. 169) that of Stomoxys calcitrans.
In 1906, a former student, Capt. R. Markham Carter,
I.M.S., forwarded to Newstead a species of tsetse-fly (namely,
G. tachinoides) from Arabia, this being the first observation of the
occurrence of Glossina outside Africa.
In 1910, Newstead described three new species of Glossina
CAnnals, Vol. IV, p. 369), namely, G. submorsitans, G. brevipalpis
and G . fuscipes. These new species were founded on an examina¬
tion of the morphological characters of the male genital armature.
A revision of Glossina, based on a study of this structure, was
made {Bull. Ent. Res., Vol. II, p. 9), and later two further new
4 o
species of this genus were described, namely, G. austeni {Annals ,
Vol.- VI, p. 129, and Bull. Ent. Res., Vol. Ill, p. 355) and
G. severitti {Annals, Vol. VII, p. 331).
In 1911, as a member of the Commission of the Royal Society
to enquire into the relation of the African fauna to human trypano¬
somiasis, Newstead proceeded to Nyasaland, and for five months
devoted himself to a study of the bionomics of the tsetse-fly
{Glossina morsitans , West.), with a view to discovering its breeding
grounds and devising means of checking its spread. The results of
this investigation were published, jointly with Dr. J. B. Davey, in
the Reports of the Commission, No. XV, p. 142. The physical
features of the country were first described and an account given
of the vegetation of the river and its borders, and the forest or fly
area. The vertebrate fauna of the district were then dealt with.
It was concluded that mpala antelopes supplied a very large propor¬
tion of the food necessary for the life and propagation of Glossina.
Two species of birds were shown to prey upon G. morsitans. The
breeding grounds of G. morsitans were thinly scattered over the
whole of the country, and large numbers did not occur in any given
spot. It was found by experiment that the average time between
each meal was about two and a half days. The period that elapsed
between the date of capture of the fly and the production of the
larvae varied from two to twenty-nine days. The duration of the
pupal period was found to be about twenty-five days. The period
of chief activity of G. morsitans was between the hours of 10 a.m.
and 4 p.m.
In 1919, Miss A. M. Evans made a study of the genital armature
of the female Glossina {Annals, Vol. XIII, p. 31). In 1918,
Newstead discovered that the innumerable papillae which form the
sculpturing on the exterior of the prominent lobes at the anal
extremity of the larvae of Glossina are respiratory openings, and
evidently function as such during the inter-uterine life of the larva.
Similar structures were found in the Hippoboscidae {Annals,
Vol. IV, p. 93).
Jointly with Carter, a new genus and three new species
of anopheline mosquitoes were described {Annals, Vol. IV,
P- 3 77 ) > namely, Dactylomyia , nov. gen., and Dactylomyia
ceylonica, Pyretophorus cardamatisi and Cellia cincta , and later
4 '
six further new species and varieties were dealt with (Annals,
Vol. V, p. 233). In 1911, Newstead and Carter founded a new
genus of Culicinae from the Amazon region, which they named
Thomasina. The type species, Thomasina longipalpis, had
previously been referred by Newstead and Thomas to the genus
Rfansonia, but it was now found that the morphological characters
of the palpi and tarsi were so markedly different from those of
Mansonia that the species could no longer remain in that genus
(Annals, Vol. IV, p. 553). Some mosquitoes of the genera
Banksinella and Taeniorhynchus were described by Carter (Annals,
Vol. VII, p. 581) with a view to establishing the affinities of certain
species and in reference also to the synonymy adopted by other
students of this group of blood-sucking insects. In 1920, Carter gave
an account of the male genital armature of the British anopheline
mosquitoes (Annals, Vol. XIII, p. 453). Extensive observa¬
tions were made by Blacklock and Carter on the bionomics of
A. plumbeus (Annals, Vol. XIII, p. 421). It was found to be
essentially a tree-hole breeder; larvae were taken from the water in
rot-holes of elm, sycamore and other trees, from 2 to 20 feet above
the ground. The breeding-places may occur in more or less isolated
trees situated sometimes within a few yards of houses. A. plumbeus
feeds on man both day and night. The observers obtained larvae
from tree-holes in December to February, and they ascertained that
thirty-five out of forty larvae survived freezing for five to thirty
minutes. (For infection experiments with A. plumbeus, see p. 12).
In 1912, Carter described three new species of the genus
Tabanus, which he named Tabanus nagamiensis, T. fulvicapillus
and T. donaldsoni, and in a subsequent study (Annals, Vol. IX,
p. 173) eight previously undescribed Tabanidae were dealt with.
He also described three new African midges (Annals, Vol. X,
p. 131), Forcipomyia lefanui, Culicoides cordiformitarsis and
Culicoides slephensi, and later (Annals, Vol. XII, p. 289) two
others, Culicoides ocrothorax and Forcipomyia ingrami, a species of
interest owing to the fact that, given favourable opportunities
for attack, its larvae prey upon the larvae of mosquitoes breeding in
rot-holes in trees. In 1920, Carter. Ingram and Macfie commenced
an exhaustive study of the Ceratopogonine midges of the Gold
Coast, with descriptions of new species. In the first account of
+2
these midges ( Annals , Vol. XIV, p. 187), after a description of the
technique employed, the observers dealt with the bionomics of the
various genera. The second instalment ( ibid., p. 211), which began
a systematic account of the Ceratopogoninae, dealt with the genus
Culicoides , and included descriptions of sixteen species, eleven of
which were new. The larvae and pupae of several species were
described in detail. The third account of this investigation ( ibid .,
p. 309) dealt with six new species belonging to three genera, of
which one of the latter is new.
In 1919, an article on the blood-sucking Nematocera was
contributed by Carter to ‘The Practice of Medicine in the Tropics *
(now in the press). This dealt with the biting flies of the
families Culicidae, Psychodidae, Chironomidae and Simultidae.
The account included general considerations regarding structure
and bionomics, the species of malaria-carrying Anopheles being
arranged in groups according to the nature of the evidence on which
they were incriminated, and the classification. In the last section,
the diagnostic characters of all the known (one hundred and twenty)
Anopheline mosquitoes were given in synoptic tables.
Newstead continued his researches on Coccidae , of which the
British species formed the subject of a monograph, in two volumes,
issued by him among the Ray Society publications, in 1900 and
1902. In 1908, he contributed an article on these insects to the
reports of the Swedish Zoological Expedition to Kilimandjaro
(Vol. II, Part 12); the same year he wrote of the scale insects and
mealy bugs of Egypt ( Liv . Univ. Inst. Comm. Res. in the Tropics
Quart. Journ., Vol. Ill, p. 14), and reported upon a collection of
Coccidae affecting plants in Java and West Africa (Journ. Econ.
Biol., Vol. Ill, p. 32). In 1910 and 1911, he described two new
species of African coccids {Journ. Econ. Biol., Vol. V, p. 18),
reported on a collection from Uganda {Bull. Ent. Res., Vol. 1,
pp. 63 and 185), on another from South and South-west Africa, and
on a third in the Berlin Zoological Museum {Mitt, aus dem Z00L
Mus. in Berl., Vol. V, p. 155). Later, he commenced a series of
studies on Coccidae, which are still being pursued {Bull. Ent. Res.,
1913-1920). In these papers, over one hundred and sixty different
species were dealt with, including the descriptions, with illustra¬
tions, of one hundred and six species and varieties new to science;
the major portion of these are serious pests to various crops under
43
cultivation in tropical and sub-tropical countries. In 1910,
Newstead served on the Special Commission appointed by the
Government of Malta to suggest means for stamping out the fluted
scale insect {I eery a pure hast), then threatening the orange-growing
industry of the island. Regulations for the suppression of the pest
were drafted and circulated.
Other work of an economic nature undertaken by Newstead
included an investigation into the food of some British birds
{Suppl. to Journ. of Board of Agric ., Vol. XV). This work, which
had extended over a period of twenty years, was based upon over
eleven hundred records, chiefly post mortem . His tentative verdict
was in favour of the birds, the records showing what an important
part is played by the majority of British birds in checking the
increase and lessening the ravages of many insect pests of plants
and crops. In 1910, Newstead dealt with some insects affecting
cultivated plants in the West Indies {Journ. Roy. Hort. Soc.,
Vol. XXXVI, p. 53), and i tx 1913, jointly with Bruce Cummings,
issued a paper on a gall-producing Psyllid from Syria {Ann. Mag.
Nat. Hist., Vol. XI, p. 306).
In 1910, Newstead went to Malta to investigate sand flies of the
genus Phlebotomus, the main object being to discover the chief breed¬
ing-places of these insects, and to recommend practical measures for
destroying them in the larval stage. It was discovered that four
distinct species of Phlebotomus occur in the island, previous investi¬
gators having notified one species only. The discovery of some
important structural details concerning the anatomy of these insects
was made and a long series of drawings illustrative of the salient
characteristics was prepared, as well as a series illustrative of the
internal anatomy {Anrtals, Vol. V, p. 139). Later, he. described
some new species {Bull. Ent. Res., Vol. Ill, p. 361, Vol. V, p. 179,
Vol. VII, p. 191, and Vol. XI, p. 305), and in 1913 dealt with three
West African species {Bull. Soe. Path. Exot ., Vol. VI, p, 124).
Besides the special studies noted above, many articles on
entomological subjects were contributed to a variety of journals by
the Department of Entomology. In addition, innumerable collec¬
tions of insects submitted by the Imperial Bureau of Entomology,
by the Belgian Government, and from other sources, were examined
and identified.
In 1915, Newstead went to France and Flanders, there to
4 +
organise measures of fly * control. A report of this work was
submitted to the War Office.
By the request of the Royal Society, Newstead and others
commenced in 1916 an investigation into the problems connected
with the damage caused to grain and flour during transit and in
storage. It was found that wheat and flour are liable to attacks
and injury by acarids, of which Aleurobius farinae was most
commonly encountered. It was established that mites will not
injure wheat and flour in which the moisture content is 11 per cent,
or under, whatever the temperature may be. The morphology and
bionomics of the infesting mites were studied, and experiments
carried out with regard to methods of destruction. Newstead and
Morris also reported upon the non-parasitic or forage acari of the
family Tyroglyphidae , to which Pillers added clinical notes derived
from veterinary experience ( Royal Society : Reports of the Grain
Pests (War) Committee , Nos. 2 and 8).
David Thomson made feeding experiments with the European
bed bug (Cimex lectularius) in various diseases (Annals, Vol. VIII,
p. 19). He found that protozoa were absent from the gut of this
species (one hundred and eighty-four examined). No acid-fast
bacilli were found in one hundred and five bed bugs fed on lepers,
nor in thirty-five others caught in bed mattresses of leper patients.
Nothing abnormal was found in bugs fed on cases of lymphadenoma,
carcinoma and malaria. Forty bugs fed on a case of spleno-
medullary leukaemia all developed numerous Charcot-Leyden
crystals in their intestines.
Dutton, Todd and Christy described the Congo floor maggot,
a blood-sucking dipterous larva found in the Congo Free State
(Memoir XIII, p. 49). The larva was stated to be semi-translucent,
of a dirty white colour, acephalous, amphipneustic, consisting of
eleven segments, and to feed mainly, or entirely, at night. The
duration of the pupal stage was a fortnight to three weeks. A light-
brown fly, caught in many huts infested with the maggot, was
subsequently identified by Austen as Auchmeromyia luteola .
Newstead (Annals, Vol. I, p. 49) noted that the true larval stage is
continued till after the formation of the puparium, and that a large
percentage of the flies escape backwards from it.
In addition to dealing with the anatomy and bionomics of
45
Ornithodoros moubata, the tick transmitting African relapsing fever
(already mentioned above), Newstead, jointly with Todd, described
a species of acarid.found infecting the lungs of monkeys, namely,
Pneumonyssus duttani (Memoir XVIII, p. 41), and later he
described another new acarid, Pneumonyssus griffithi, found in the
lungs of the Rhesus monkey ( ibid ., p. 47).
PROTOZOOLOGY
Dutton, Todd and Tobey gave an account of certain parasitic
protozoa observed by them in Africa (Memoir XXI, p. 87, and
Annals , Vol. I, p. 285) in mammals, birds and reptiles, including
full descriptions and figures of some forms of Leucocytozoon
ziemanni, parasitic in a grey hawk of the Congo. Fantham studied
the leucocytozoon, L. lovat't, of the red grouse, La go pus scoticus t
and observed the occurrence of schizogony in its life cycle (Annals,
Vol. IV, p. 255). He also studied a flagellate found in the
alimentary tract of the body louse, which he named Herpetomonas
pediculi (.Annals , Vol. VI, p. 25), and of which he demonstrated
the complete life cycle, notifying its occurrence in lice in England,
as well as in India and Tunisia (ibid., p. 403). Another Herpeto-
monas, H. slratiomyia, sp. n., was discovered by Fantham and
Porter (Annals, Vol. VII, p. 609) parasitic on the larvae, pupae and
imagines of the flies Stratiomyia chameleon and S. pot amid a.
Research into induced herpetomoniasis in birds resulted in
producing this condition in canaries, sparrows and martins by
feeding them on insects containing herpetomonads; in some cases
the infection was fatal. It was found that the cycle of the
flagellate in the avian host resembled morphologically that in the
insect (Annals, Vol. IX, p. 543). Fantham and Porter studied the
effects on their hosts of certain Myxosporidia inhabiting the gall
bladders of various fish (Annals, Vol. VI, p. 467). In 1916,
Fantham contributed the section on the Protozoa to ‘The Animal
Parasites of Man/ issued jointly with Stephens and Theobald.
Seidelin described some blood parasites in reptiles (Annals,
Vol. V, p. 371), and also some species of Klossiella in the kidney
of a guinea-pig (Annals, Vol. VIII, p. 553). E. H. Ross observed
the development of a leucocytozoon in a guinea-pig, for which he
4 6
proposed the name Lymphocytozoon cobayae ([Proc . Roy . Soc., B.,
Vol. LXXX, p. 67); Sinton prosecuted research into the morphology
and biology of Prowazekia urinaria {Annals , Vol. VI, p. 245); and
O'Farrell, in a study of hereditary infection, with special reference
to its occurrence in Hyalomma aegyptium infected with Crithidia
hyalontmae , gave an account of the four periods in the life cycle of
this flagellate {Annals, Vol. VII, p. 545).
In 1917, Smith and Matthews investigated the incidence of
intestinal protozoa in two hundred and fifty patients admitted to
hospital for diseases other than dysentery {Annals, Vol. X, p. 361).
Entamoeba histolytica was found in 8 per cent, of the cases, E. coli
in 19*2 per cent., G . intestinalis in 8 per cent., C. mesnili in
2 per cent., and T. intestinalis in 1*7 per cent. Of the two hundred
and fifty cases examined, two hundred and two were suffering from
non-intestinal complaints, and of this number £*4 per cent, were
found to be harbouring cysts of E . histolytica . Among ninety-one
men who had been to France only, two were discovered to be
‘carriers 1 of E. histolytica . In a further investigation {Annals,
Vol. XI, p. 183), two hundred non-dysenteric patients were
examined, and protozoal infections found in 34*5 per cent.;
E . histolytica in 7*5 per cent. Matthews described and figured the
characteristic morphological features of cysts of the common
intestinal protozoa of man {Annals, Vol. XII, p. 17), and later
made a mensurative study of the cysts of E . coli {Annals, Vol. XII,
p. 259), and traced the course and duration of an infection with
this parasite {Annals, Vol. XIII, p. 17). Matthews and Smith
investigated the spread and incidence of intestinal protozoal infec¬
tions in the population of Great Britain. The first selected
population consisted of four hundred and fifty civilians in the
Liverpool Royal Infirmary {Annals, Vol. XII, p. 349), of which
i‘5 per cent, were found to harbour E. histolytica and 6*7 per cent.
E . coli. The figures for army recruits, of which one thousand and
ninety-eight cases were examined, were E. histolytica 5*6 per cent.,
E . coli 18*2 per cent., E. nana 2*4 per cent., G . intestinalis
6*o per cent., and C. mesnili two cases. In five hundred and forty-
eight children, all under the age of twelve {Annals, Vol. XII,
p. 361), G. intestinalis was the parasite most commonly found. Of
two hundred and seven male asylum patients {Annals, Vol. XIII,
47
p. 91), 9‘7 per cent, were found to be infected with E. histolytica ,
45’9 per cent, with E . coli , and 23*2 per cent, with C. mesnili.
University and School cadets were also • examined; the same
protozoa were found as amongst other series, but the number of
cases recorded was too small to allow of conclusions being, drawn
as to incidence amongst this higher social class.
In 1906, Drs. Fantham and Porter investigated the Isle of Wight
bee disease ( Annals , Vol. VI, p. 163). It was found that the
disease was due to a minute microsporidian parasite, Nosema apis,
sp. n., which gained access by the mouth to the intestines. Experi¬
mental work proving the pathogenicity of the parasite was carried
out. It was found that 'Nosema apis was harboured by other insects
besides the hive bees ( Annals , Vol. VII, p. 569). It was considered
that a bee, itself apparently immune, can be a parasite carrier. A
morphological study of Nosema apis was made, and the two phases
in its life cycle demonstrated: (1) a multiplicative phase, termed
merogony, which occurs in the epithelium of the chyle stomach
and intestines of the bee; (2) a second phase, termed sporogony,
leading to the formation of minute, resistant resting spores, which
are shed in the faeces of the bee, fouling the surroundings of the
hive and producing.infection of fresh bees when swallowed in food
or drink. An allied organism, Nosema bombi , sp. n., parasitic in,
and pathogenic to, bumble bees, was discovered, and its life cycle,
and suggested economic measures of control, set forth in a subsequent
paper {Annals, Vol. VIII, p. 623).
49
ON THE ‘ ARNETH COUNT ’ IN HOOK¬
WORM-INFECTED WHITE CHILDREN
IN NORTH QUEENSLAND
BY
A. BREINL
From the Australian Institute of Tropical Medicine ,
Townsville
(Received for publication 31 January , 1921)
In the past, work has been carried out by Breinl and Priestly
(1914) on Ameth counts of healthy white school children, who had
spent their lives in the tropical parts of North Queensland. The
observations proved a decided shift of the Ameth index to the left,
when compared with that of normal individuals in Europe. Later,
this investigation was extended to healthy aboriginal children in
Northern Australia (1917), and, furthermore, to native children in
New Guinea (1915), living in an area where malaria, yaws and
other parasitic diseases were found to be endemic.
A shift of the Ameth index to the left was found in healthy
aboriginal children of North Queensland analogous to that of
children of European descent in North Queensland, but a much more
pronounced shift was found in the native children of New Guinea.
Taking advantage of recent opportunities, blood smears
obtained from white children in North Queensland, suffering from
ancylostomiasis were examined in order to determine the Ameth
index. A number of the smears were collected by one of the
members of the staff of the hookworm campaign, carrying out work
at present in North Queensland; others were obtained from children
of various ages, who underwent treatment for hookworm infection
in the Townsville Hospital.
Previous observations by Knapp (1915) in India showed in the
blood of hookworm patients a distinct shift of the Ameth index to
So
the right, but the results, according to his own statement, ‘were on
the whole equivocal,* and he proposed to carry out further research.
Macfie (1916), working on the Gold Coast, confirmed to a
certain extent Knapp’s tentative results. Out of seventeen counts
made on hookworm patients, about 30 per cent, showed, when
compared with those of healthy natives, an actual shift to the right,
41 per cent, had a relative shift to the right, that is, 1 a slighter
degree of shift to the left than is found in apparently healthy
natives,* and 29 per cent, had a definite shift to the left. He
concluded from his results that ‘there appeared unquestionably a
tendency to develop a shift to the right in patients infected with
hookworms.*
In the present investigation the same technique was employed as
in the previous work; all counts were performed by myself.
Dr. Priestly and myself having performed the previous counts. In
this way the results, so far as technique is concerned, are comparable
with those obtained previously, and the personal source of error has
been, so far as possible, excluded.
Two hundred consecutive leucocytes were counted in two sets of
100, and only when the two sets of figures differed but slightly
were the counts taken into consideration.
All the children from whom the blood was obtained lived in
areas where malaria is practically unknown, and any definite change
found must be attributed to the effects of hookworm infection.
The figures (Table I) were separated according to age groups,
and the table shows that the averages for the age groups
between five and fifteen years are fairly constant. The Ameth
index for children two and three years old is much lower, and
approaches that found in healthy North Queensland children. The
disproportionate rise for children four years of age may be due to
the small number of observations. The total averages prove
conclusively that the average Arneth index in hookworm-infected
children shows a decided shift to the left.
A comparison of the Ameth index of hookworm-infected white
children in North Queensland with that of children living in New
Guinea shows a striking similarity, and strengthens the assumption
that the comparative increase in the Ameth index in the latter
locality was due to the great incidence of latent and active infection
Arnith Clashcation Differential Counts
51
52
amongst children, amongst whom hookworm infection is probably
widespread.
The significance of the shift of the Ameth index to the left is
still uncertain. It is, however, possible that in such diseases as
hookworm infection and malaria, where the destruction of red cells
goes, for a time at least, hand in hand with an increased activity
of the blood-forming organs, the increased activity extends to the
new formation of leucocytes, and in consequence a greater number of
young leucocytes are met with in the peripheral blood.
The differential counts indicate that in younger children—
between two and four years of age—the relative number of
polymorphonuclear neutrophile leucocytes is decreased, whereas that
of the lymphocytes is increased.
After the seventh year the relative proportion of the various forms
of white blood corpuscles is fairly constant.
As is to be expected, the relative increase in the number of
eosinophile leucocytes is well pronounced throughout our series of
counts.
REFERENCES
Bizutl and Pkiutly (1914). Ann. Trap, Med, and Porotit.y VoL VIII, p. $6$.
-(1915). Aum. Trop. Med. and Porosity VoL IX, p. 495
- ( , 9 I 7 )* Trap. Med. and Parasit., VoL X, p. 427.
Knapp (1915). Indian Med. Gan ., VoL L, p. 65.
Macfik (1917). Report of the Accra Laboratory for 1916, p. 44. Churchill, London.
53
BRONCHOMONILIASIS COMPLICATING
PULMONARY TUBERCULOSIS IN A
NATIVE OF THE GOLD COAST,
WEST AFRICA
BY
J. W. S. MACFIE
AND
A. INGRAM
(Received for publication 2 February, 1921)
Since Castellani discovered the condition in Ceylon in 1905,
bronchomoniliasis has been identified in many parts of the world,
especially in tropical and sub-tropical countries. In Africa,
Chalmers and Macdonald (1920) studied a number of cases in the
Sudan and Egypt, and Pijper (1917) has noted the presence of the
disease in South Africa, but, so far as we are able to ascertain, no
cases have hitherto been recorded from West Africa. For this
reason, a short account will be given of a case which has recently
come under our notice at Accra, in the Gold Coast, West Africa.
The case was not a pure bronchomoniliasis, but occurred in a
patient 'suffering from pulmonary tuberculosis, a form of mixed
infection which apparently is not uncommon, and has been observed
previously by de Mello and Fernandes, Castellani and Chalmers,
and others.
History. We are indebted to Dr. J. R. Moffatt for the following
history of the case. I. D., a native (Buzaburime), aged about
twenty-five years and a member of the Police Force, admitted to the
Native Hospital, Accra, on the 29th of September, 1920. The
patient stated that he had suffered severely from cough for at least
two months previous to admission. Upon being closely questioned,
he further admitted that for at least eighteen months he had
experienced attacks of illness, accompanied by cough, at irregular
intervals. Physical examination revealed dullness at the apex of
the right lung, chiefly supra-davicular; at the base of the left lung
there was a considerable area of dullness extending as high as the
54
angle of the scapula, and at its upper portion, especially on the
anterior aspect, bounded by a hyper-resonant area. The cough
was frequent and harrassing; the sputum copious and in appearance
like thin flour paste. Sweating was inconsiderable. Although the
breathing was rapid, the patient never suffered from dyspnoea. The
temperature chart kept whilst the patient was in hospital showed
an irregular fever similar to that which might hcLve been expected in
a case of pulmonary tuberculosis. The treatment given was cod-
liver oil and, for a few days only, potassium iodide. During the
last two weeks of his illness the patient showed some improvement
and gained weight; on the night of the 8th of November, however,
he had a sudden and copious haemoptysis and died within an hour.
A specimen of the sputum of this case was forwarded to the
laboratory for examination as to the presence of tubercle bacilli upon
the 30th of September; none were found on this occasion, but it was
noted that the sputum had a curious appearance, suggestive of saliva
containing small particles of macerated bread, and accordingly
another specimen was asked for which should be taken after
thoroughly washing the mouth with a weak antiseptic solution. On
the 2nd of October the second specimen of the sputum was examined,
and was found to contain numerous yeast-like cells {Monilia sp.)
but no tubercle bacilli. Cultures were made from this specimen of
the sputum upon Sabouraud’s maltose agar and glucose agar, and
within twenty-four hours a copious creamy-white growth of the
Monilia had made its appearance on both media. On the 5th of
October the sputum again showed yeast-like Monilia cells and also
a few coarse hyphae; tubercle bacilli were not detected. As
potassium iodide has proved a valuable remedy in broncho-
moniliasis, it was suggested that it should be tried in this case, and
this was done for a few days, ten grains being given thrice daily.
The effect was to reduce the number of Monilia cells in the sputum
very greatly, and to reveal the presence of tubercle bacilli which in
all probability had been overlooked at previous examinations. A
specimen of the sputum examined on the 8th of October showed
very numerous tubercle bacilli and no Monilia cells, and as the
potassium iodide appeared to be causing^ the patient some
discomfort, owing to the increase in the quantity of his sputum, it
was then stopped. Sputum examined on the 14th of October, and
55
later, showed that the Monilia cells had reappeared and that the
number of tubercle bacilli appeared to be fewer. The patient
died on the 8th of November as the result of an haemoptysis, and a \
post-mortem examination of the body was made on the following !
morning.
Post-mortem Examination. The following were the notes made
at the examination. Body: that of a young native man, rather
emaciated. Abdominal cavity and its contents: appeared to be
normal. Spleen : not enlarged, weight seven and a half ounces, no
visible morbid condition.' Kidneys and liver: showed venous
engorgement but no other pathological signs. Gall bladder:
collapsed, empty. Mesenteric glands : not enlarged. Right lung :
adherent to the chest wall at its apex; on removal and section a
cavity about the size of a walnut was found at the apex, it contained
a blood clot; the whole of the upper lobe and a portion of the middle
studded with small tubercles. Left lung: completely collapsed,
visceral and parietal pleurae very greatly thickened and of a creamy-
yellow colour; the pleura covered in parts with a deposit of the
colour and consistence of cream cheese, the pleural cavity contained
about four ounces of turbid Straw-coloured fluid; the substance of
the lung studded with numerous caseating tubercular nodules, and
at one point apparently communicating with the pleural cavity.
Lymphatic glands at the root of the neck and in the mediastina:
enlarged, tuberculous.
Smears of the creamy exudate in the left pleural cavity showed
numerous tubercle bacilli and yeast-like Monilia cells, also a
considerable number of short septate branching hyphae. Sections
of the lungs showed that both were the seat of advanced tubercular
disease; sections of the thickened pleura of the left lung and of a
mass of the pleural exudate showed also the presence of Monilia.
Organism. The Monilia found in this case was easily obtained
in pure culture by inoculating tubes of Sabouraud’s maltose agar
and glucose agar. It was gram-positive but not acid fast. It grew
well on most solid media, but especially well on glucose agar,
and produced rapidly a diffuse, spreading, creamy-white growth.
Under anaerobic conditions its growth was slower. Gelatin and
blood serum were not liquified by it, and did not become pigmented.
In broth and peptone water it caused a white deposit to be thrown
56
down whilst the media themselves remained clear; m peptone
water a slight surface pellicle was formed. It produced a thick
white growth on potato. On solid media the growth was almost
entirely composed of yeast-like cells; in some fluid media hyphae
predominated.
Its qualitative bio-chemical
follows: —
Arabinose .
... 0
Rhamnose (isodulcite)
... 0
Galactose .
... AG
Glucose.
... AG
Laevulose .
... AGs
Mannose .
... O
Lactose.
... O
Maltose.
... AGs
Saccharose .
... AG
Amylum .
... O
Dextrin.
... O
Glycogen .
... O
reactions may be tabulated as
Inulin
Amygdalin
Helicift ...
Phlorrhizin
Salicin ...
Glycerol...
Erythrol ...
Adonitol
Dulcitol ...
I nosite ...
Mannitol...
Sorbitol ...
The symbols representing : A—acid; G—gas; s—slight; and
O—neither acid nor gas.
The production of gas in laevulose and maltose was slight. If
the cultures were kept for two weeks or longer the acidity produced
in the five sugary media indicated tended to be superseded by
alkalinity; this was earliest seen and most pronounced in glucose
and saccharose. At first no change was produced in litmus milk,
but later, after about ten days, alkalinity developed; no dot was
formed and the medium was neither decolourised nor peptonised.
Indol was not produced in peptone water.
As gas was produced in glucose, laevulose, maltose, galactose,
and saccharose, the organism comes into the fifth group of species
of Monilia , called the Tropicalis group , according to the classifica¬
tion of Castellani and Chalmers (1919). In this group are placed
(loc . cit . p. 1084) M. tropicalis , Cast., M . paratropicalis , Cast.,
M. pulmonalis , Cast., M . nivea , Cast., M . insolita , Cast., and
M. enterica , Cast.; but from the table given by the same authors
(pp. 1082-1083) it would appear that M. faecalis , Cast., and
M . metatropicalis , Cast., should also be included. A somewhat
later table given by Castellani (1920) differs slightly from that given
by Castellani and Chalmers and omits certain species whilst intro-
000000000000
57
during some additional ones. According to it, acid and gas are
produced in the five sugary media mentioned by five species only,
namely, M . enterica, M. faecalis , M. metatropicalis , M. para-
tropicalis, and M. tropicalis .
Reverting to the species given by Castellani and Chalmers,
which include the five given by Castellani alone and three others,
it will be seen that the bio-chemical reactions of the organism isolated
from our case do not agree entirely with those of any of them (see
table). The reaction in litmus milk suffices to distinguish it from
Table showing the more important biochemical reactions of the species of Monilia of the Tropicalis group .
Spedes of Monilia
Litmus
milk
Glucose
Laevulose
Maltose
Galactose
Saccharose
Mannite
Dextrin
1
Broth
Mm enterica ..
°/
/Aik
AG
AG
AG
AG
AG
As
As
.
o
0
C
At. faecalis .
A /
/DP«
AG
AG
AG
AGs
AGs
O
O
o
o
C
M. insolita .
As j
/ Aik
AG
AG
AG
AG
AG
As
O
o
o
C
M. metatropicalis
AC
AG
AG
AG
AG
AG
O
o
o
o
C
At. nivea ..
°l
/Aik
AG
AG
AG
AG
AGs
O
o
AG
o
C
M. paratropicalis
A 7au
AG
AG
AG
AG
AG
o
Avs
O
o
CTP
M . pulmonalis
°/ AlkD
AG
AG
AG
AGs
AG
Avs
o
A
AGs
CTP
At, tropicalis .
A
i
AG
AG
AG
AGs
AGs
O
O
O
o
C
Species isolated from case of
Bronchomoniliasis at
Accra
0 /
'Aik
AG
AGs
AGs
AG
AG
O
O
?
o
C
A = acid : G = gas j s * slight * vs = very slight; O neither add nor gas produced ; Aik = alkalinity;
C — clot; D — decolonisation ; C = clear ; jjCTP — clear thin pellicle ; P — peptonisation.
all the other species of the Tropicalis group excepting M . enterica
and M. tiivea . The former of these two produces slight acidity in
mannitol and dextrin, reactions which are not produced by our
species. As regards the latter, M. tiivea , acid and gas are produced
58
in raffinose, and acid but only a small amount of gas in saccharose.
We w,ere unable to test the reaction of our species in raffinose, but
in saccharose much gas was produced, and only a small amount in
laevulose and maltose. It is admitted, however, that many of the
species of the Genus Monilia have not permanent bio-chemical
reactions, a point emphasised by Castellani himself, and if they are
liable to vary outside the body, it seems not unlikely that they may
vary also according to their host. The very slight differences noted
between the bio-chemical reactions of the organism recently isolated
by us and those of M. nivea are, therefore, probably unimportant.
M. nivea was originally found in sputum, and is considered by
Castellani and Chalmers to be of doubtful pathogenicity. It is of
interest, therefore, to recall that the organism resembling this species
which we have isolated was found not only in the sputum but also,
after death, in the body of the patient.
SUMMARY
A case is recorded in which bronchomoniliasis complicated
pulmonary tuberculosis in a native of the Gold Coast at Accra.
The patient died of an haemoptysis whilst under observation. At
the post-mortem examination both lungs were found to be
tuberculous. The left lung was collapsed, and the pleural cavity
partially filled with exudate. In this exudate and in the thickened
pleura over the lung Monilia was present.
The organism, which belonged to the Tropicalis group of
Castellani and Chalmers, closely resembled in bio-chemical reactions
M. nivea , Cast. (1910); without raffinose we are unable to state
whether the species found at Accra is distinct from M. nivea, Cast.
REFERENCES
Castellani, A. (1905). Ceylon Medical Reports.
- (1920). Milroy Lectures on the Higher Fungi. Journ. Trop. Med. and Hyp., Vol.
XXIII, p. 118.
- and Chalmers, A. J. (1919). Manual of Tropical Medicine , Third Edition, pp. 1070-
1092. London : Bailliere, Tindall and Cox.
Chalmers, A. J., and Macdonald, N. (1920). Bronchomoniliasis in the Anglo-Egyptian
Sudan and Egypt. Journ. Trop. Med. and Hyp., Vol. XXIII, pp. 1-7.
de Mello, F., and Fernandes, L. G. (Reviewed in the Trop. Dt*. Bull ., Vol. XIV, p. 244).
Pijper, A. (1917). Med. Journ. S. Africa, Vol. XII, pp. 129-130.
59
NOTES ON A CASE OF INDIGENOUS
INFECTION WITH P. FALCIPARUM
BY
B. BLACKLOCK
(Received for publication 8 February , 1921)
In their preliminary note on this case, Glynn and Matthews
(1920) have furnished an account of the findings at the post-mortem
examination and also details of the history. For convenience we
may recapitulate here briefly the main facts.
History . The patient was a girl, aged 18, who had never been
out of the British Isles; she was born in Liverpool and had once
been South—to London in 1914, from July to September, during
which period she spent three weeks at Littlehampton; she had lived
for the last ten years in the same house in Liverpool. She died on
October 12th, 1920, after a period of illness which commenced (on
her return from a holiday in a northern health resort) with
4 feverishness and flushed appearance* on October 1st, followed on
October 2nd and 3rd and several other occasions by vomiting. On
October 8th she was overcome with faintness in the street, and had
to be helped home. Her condition rapidly became worse with
severe headache, thirst, photophobia, delirium, paresis of legs,
anuria, and finally coma. The temperature on October nth, the
day before.death, was ioo°F., and on October 12th, just before
death, io2 °F. Further investigation elicited the facts that she had
always been pale, had suffered from headaches since Christmas,
1919, and that she had vomited on September 29th, 1920.
The following notes on the post-mortem examination are quoted
from the paper mentioned above: —
‘ The post-mortem examination was made twenty hours after death. Blood :
anaemic. Brain : apparently normal, no meningitis. Lungs : a moderate amount
of oedema and muco-purulent bronchitis. Heart : weight, 8 07.., normal, no fatty
degeneration. Mouth : healthy, no trace of pyorrhoea. Oesophagus, stomach
and intestines opened from end to end—normal, no evidence of blood. Some of
the lumbar lymphatic glands slightly enlarged, and red like haemolymph glands.
Liver : weight, 3 lbs., slightly browner than normal, no haemosiderin reaction.
Spleen : I lb. 7 oz., retains its shape, of normal consistency, but dark, rather like
6o
malaria. Pancreas, suprarenals, kidneys and bladder normal. Uterus: normal
size, but menstruating ; one haemorrhagic corpus luteum. Bone marrow of the
shaft of the femur a uniform terra-cotta red, that of the sternum redder a nd more
succulent than normal/
The following is a short account of my observations on the
material examined. It is convenient to state here that the specimens
from this case, which were demonstrated by me at a meeting of the
Royal Society of Tropical Medicine and Hygiene, were by some
inadvertence included under Col. James’s exhibits, and erroneously
stated to have been derived from a case in Sheemess. (See Lancet ,
January 1st, p. 26.)
Blood film . About a third of the red corpuscles contained young
trophozoites, but owing probably to post-mortem changes the
parasites appeared small and contracted, giving an appearance
similar to that seen in sections. Not only was the total number of
red cells infected large, but also multiple infection was common,
many cells containing two, three, four, up to five parasites.
Gametocytes were present in small numbers most of them being
fully developed crescents, some of them presenting a peculiar
appearance owing to the red cell being visible all round the
parasite and extending some distance beyond it, and not chiefly
in the concavity of the crescent; other forms were oval or
spherical, and in addition there were seen solid-looking parasites
of irregular outline which might be interpreted as developing forms
of gametocyte. A small number of segmenting forms with a number
of merozoites varying from ten to twenty-four also occurred in the
blood, the crescents and fully divided forms being in about equal
proportion.
Nucleated red cells were numerous, as was observed in the
preliminary note, the figure given being 3*8 nucleated reds to 100
leucocytes (500 counted), while my figure is slightly higher, 4*9 to
100 leucocytes (1,000 counted). They presented much diversity in
the form of the nucleus, single-nucleated forms predominating, but
forms with double and multipartite nuclei also being found.
Pigment was present in mononuclear leucocytes and also free in the
plasma, some in the latter possibly being due to the breaking of
segmenting forms in making the film.
Spleen smear. The chief feature to attract attention was the
occurrence of very numerous segmenting forms, not all at the same
stage of sub-division, the merozoites numbering from eight to
twenty-two. There were a few crescentic, oval, and spherical
gametocytes. An occasional parasite, even in the segmenting stage,
was found ingested by phagocytes. Pigment, both black and
golden yellow, was present in considerable amount, ingested by
large mononuclear white cells.
Bone marrow smear . Here crescentic and oval forms of gameto-
cyte were numerous in quite different proportions to the numbers in
either the blood or spleen smears. A few segmenting forms occurred
and many small trophozoites, of which some were free. Among the
large mononuclear cells, many were observed which contained
eosinophil granules, and many also containing pigment.
Several questions of interest arise in connection with this case,
and it is necessary, in spite of the limited character of the observa¬
tions made on the case during the period of her illness and the
small amount of material available for examination, to endeavour to
answer some of the questions.
1. Was this a primary acute attack following directly the
incubation period, or a secondary acute attack supervening in a
person already infected for a considerable time ?
2. By what means did the patient acquire infection?
3. At what time and where did she become infected ?
I. WAS THIS A PRIMARY OR SECONDARY ACUTE ATTACK ?
(a) Primary acute attack.
The absence of any history, previous to October 1st, 1920, ot
fever, shivering or sweating, is in favour of this being a primary
acute attack.
Spleen . Support is also lent to this theory by certain appear¬
ances presented by the spleen. The noteworthy condition seen in
section of this organ is the remarkable congestion, with dilatation
of the sinuses. These facts are illustrated by photograph I, repre¬
senting a section from the spleen of the case under discussion. That
the large size of the spleen is due to a recent enlargement of the
organ is further demonstrated when we observe that the increase
of size is due to engorgement with blood and not to increase of
fibrotic tissue, nor to the cellular elements of the spleen itself.
(£) Secondary acute attack.
It is quite possible for a person to harbour and present for
considerable periods in his blood P. falciparum without the produc¬
tion of definite signs and symptoms, provided the person is taking
quinine or other drugs. This fact has been observed frequently,
and instances will be found in the work of Stephens and his
collaborators (1917). Similarly it is recognised that persons
suffering from relapses of malignant tertian malaria may have
parasites present in the peripheral blood for varying periods before
Photo. I. Section of the spleen showing the Photo. II. Section of normal spleen.
Malpighian body with vessel cut obliquely and X about 280.
below it the accumulation of red blood corpuscles. Photographs lent by Pro fessor Gly
X about 280.
the next attack occurs. But it is also possible that a person who is
infected may, even if untreated, remain without definite signs or
symptoms for a long period and then suddenly develop an acute
attack; instances of incubation periods up to ‘months’ are
mentioned by Craig (1914). It should be observed, however, that
in experimental infections in which healthy persons are injected with
blood containing parasites or are bitten by anophelines containing
sporozoites in their salivary glands, such long incubation periods
are not recorded. Ross, whose contribution to the elucidation of
63
this case is dealt with more fully below, does not supply information
which is of assistance concerning this point. In the case under
consideration the only evidence of old standing symptoms or signs
which might possibly be attributed to malaria are that the girl had
suffered from headaches from Christmas, 1919, her pallor, and the
fact that she vomited on September 29th. With regard to these
phenomena, they may be attributed to so many other causes that
they can give us no clue.
Spleen . The features which may be regarded as indicative of
chronicity are the size (644 grms.) and lack of diffluence. As
regards the former, Dudgeon and Clarke (1918-19) give the weight
of the spleen in a series of fatal cases of pernicious malaria due to
P. falciparum as varying from 250 to 450 grms.; the highest weight
in a chronic case was 960 grms. On the other hand, James (1920)
emphasises the great splenic enlargement observed in indigenous
cases of simple tertian malaria in children. Previously Osier (1901)
mentioned a case of death from accident during the second week
of simple tertian malaria where the spleen weighed 800 grms., was
very dark red, and diffluent.
II. BY WHAT MEANS DID THE PATIENT ACQUIRE
INFECTION ?
The two known methods by which it is possible to transmit
malaria to a healthy person are the inoculation of infected blood and
the transmission by the bite of a mosquito with infected salivary
glands. The incubation period, using such methods, varies
considerably for malignant tertian parasites, and it is necessary to
enquire into the periods and how they are estimated.
The inoculation of Infected Blood into Healthy Persons
Of the numerous records obtained by inoculating blood containing
malaria parasites into healthy persons only a small number is
concerned with the malignant tertian parasite. Some of the early
experiments collected by Thayer and Hewetson (1895) are, for
comparative purposes, set out here in tabular form.
It will be seen from Table I that in the early experiments the
parasite incubation was long. These early experiments are, however,
6 4
open to several objections. In Gualdi and Antolisei's first two
experiments the inoculation of blood from quartan patients resulted
in the appearance, in the subjects of inoculation, of malignant
tertian parasites. The authors had to account for this by discovering
later that the patient whose blood was inoculated harboured both
forms of parasite. In the third case recorded by Gualdi and
Antolisei, and that of Di Mattei, crescents only were visible in the
blood injected, while in the last case, that of Sacharow, not only
was the injection small in amount and given subcutaneously, but also
the blood was taken from leeches which had been fed four days
previously on the infected person and which, since feeding, had
been kept on ice.
Table I.
Plasmodium falciparum. Incubation period following blood inoculation. Early experiments.
Observer
1
Route of
Injection
j
Amount
of Blood
injected
c.c.
Day on which,
in inoculated person
Remarks
Fever
commenced
Parasites
appeared
Gualdi and Antolisei
1 i
Intravenous ...
i
3
'
IO
IO
!
Source of blood, quartan cases.
Gualdi and Antolisei
1 Intravenous ...
i
3
12
12
Source of blood, quartan cases.
Gualdi and Antolisei
1 ?
1
1
2
9
IO
Crescents only found in blood
injected. Crescents seen in
inoculated person in i& days.
Di Mattei .
Intravenous ...
1
?
16
In a few
days after
injection
Crescents only found in blood
injected. Crescents seen in
inoculated person in 25 days.
Sacharow .
Subcutaneous
i
I
|
12
13
Blood kept in leeches on ice
4 days before injection.
Experiments were performed later by Bastianelli and Bignami,
which are set out in Table II, and from this series a very different
idea of the incubation period following inoculation is obtained.
Parasites appeared in the peripheral blood in so short a period as
two days after injection of 2 c.c. of infected blood, and infection
resulted after such small quantities as 0*2 c.c. of blood injected.
No mention is made as to the date of crescents appearing in the
blood in these cases.
65
It appears, then, that the incubation period after experimental
inoculation, whether as regards occurrence of fever or appearance of
parasites in the blood/ may be exceedingly brief in malignant
tertian malaria.
Ross (1920) has suggested the possibility of this case having
acquired infection by inoculation of blood. There was no evidence
in the history of the case that such a mode of infection could have
been the cause. She had cocaine injections for removal of teeth in
July, 1919, and in February, 1920; for the removal of the teeth,
mentioned as done on 14th September, 1920, the anaesthetic used
was ether, and no evidence was obtained either that any such
injections as morphia had been given, or that infection could have
Tabu II.
Plasmodium Jalciparum. Incubation period following blood inoculation. Later experiments.
Observer
Route of
Injection
Amount
of blood
injected
c.c.
Day on which,
in inoculated person
Remarks
Fever
commenced
Parasites
appeared
Bastiauelli and
Bignami
Intravenous ...
1
2*0
2
2
Few parasites seen in injected
blood.
...
S-o
2
2
...
°75
5
?
Few parasites seen in injected
blood.
1
1
...
0*2
4
?
Fair number of parasites seen in
injected blood.
arisen from abrasions of skin or mucous membranes having
facilitated transmission. It is regrettable that Ross, who has
sources of information inaccessible to others, should not have
communicated the facts in such a form as to have added to our
knowledge of the various points of interest which must have arisen
as the result of his observations. The incubation period of
malignant tertian malaria under such circumstances, the dates of
the first appearance of symptoms and parasites, and also the
important question as to the time of the appearance of crescents, are
all matters of sufficient interest to make welcome any additional
facts relating to them.
66
Incubation period of P. FALCIPARUM following the Bites of Infected
Anopheline Mosquitoes
#
The records of experiments of this nature do not comprise
numerous observations, and in some cases the data given are
insufficient for our present purpose. In Table III are shown the
results of some experiments in which more complete data are given
by the observers.
Table III.
Plasmodium falciparum , incubation period following the bites of anophelines previously infected.
Healthy individual
Tempera-
Observer
Anopheline
used
or
Bred
Number
of mos¬
quitoes
fed
Number
of days
they were
fed
Fever
com¬
menced
in days
Parasites
appeared
in days
< at which
| mos-
' quitoes
j ke P‘
Remarks
Bastianelli and
Bignami (1899)
A. maculipennis ...
;
Wild ...
2
9-12
i
1
10-13
?o°C.
(
The person on vrhsa
the mosquitoesven
infected develops
Simple Tertian
Malaria in 17-19
days.
Schtlffner (1901) ...
A. vagus .
Wild ...
?
2
15-16
17-18
Room
Large rings found.
Jancs6 (1908)
A . maculipennis ...
Wdd ...
2
1
1
•
1
12
12
24 0 C.
1
Quinine. 1 gm. on ~t
day after bites.
Jancs<S (1908)
A, maculipennis ...
Wild ...
I
2
1
1
10-15
11-16
| 20° C.
1
Quinine given afte
first bite, 1 gm.
1 gm., 1*5 gm.
x 5 gm., on
9th, 10th. 1 ith day
respectively.
Mitfmain ^1916) •••
A. quadrimaculatus
Bred ...
I
1
10
1 IO
21-22° C.
Accidental infection.
1
In these experiments the maximum parasitic incubation period
of P. falciparum after infection by means of bites of infective
anophelines is eighteen days, the minimum ten days.
Before accepting the results of these observers, however, it is
necessary to enquire more closely whether they are justified in
regarding the experiments as conclusive. In Bastianelli and
Bignami’s experiment wild A. maculipennis were fed, for the
purpose of infecting them, on a patient suffering from malignant
tertian fever; this patient after seventeen to nineteen days from the
commencement of feeding the mosquitoes on him showed in his blood
the parasites of simple tertian fever. If the wild mosquitoes
67
infected him with Plasmodium vivax it appears not improbable that
they might also infect with P. vivax the healthy person on whom
they fed, and, therefore, one would expect that corroborative
evidence such as the date of appearance of crescents in this case
should be given. Apart from this, there is evidence, in some of
these records, that the population used for experiment was subject
to multiple infection. Reference has already been made to the fact
that Gualdi and Antolisei, using the blood of patients supposed to
be carrying P. malariae , obtained in the person injected
P . falciparum This may be capable of many explanations, even
excluding Laveran’s theory, recently revived by Grassi (1920), of
the unicity of the parasite, for example that they conveyed the
parasite of malignant tertian fever by the inoculation, or that the
person inoculated was himself already suffering from infection with
P. falciparum or that he acquired such infection from mosquitoes
independently of the inoculation. In the same way, in Bastianelli
and Bignami’s case, additional evidence as regards the species of
parasite recovered is not available. In Jancs6’s experiments the
administration of quinine may have influenced the incubation
perod. Mitzmain's accidental case appears to be the most satis¬
factory.
These considerations are mentioned in order to draw attention
to the limited amount of reliable information available concerning
the incubation period of P. falciparum . Factors such as the use of
wild mosquitoes, experimenting among an already infected popula¬
tion, and the administratipn of quinine after the infective bite reduce
materially the value to be attached to the figures. A further point
to be mentioned, which may seriously affect the parasitic incubation
periods given above, is that, as noticed previously, parasites are
frequently present in relapse cases before a temperature reaction
occurs. The importance of remembering this fact is seen when we
consider Tables I, II and III. It will be noted that in these experi¬
ments, fourteen in number, parasites were always discovered either
after the day of fever or on the day of fever, with the exception of
one case (di Mgttei), in which they were found ‘ a few days after
injection/
From what has been said, therefore, it is seen that the incubation
period of malignant tertian malaria is a subject which would repay
68
further study. At present the evidence points to a minimum
incubation, for inoculation of two days, and for mosquito bites of
ten days. In view of the absence of evidence of inoculation infec¬
tion in this case, we must suppose that an anopheline was the means
of infection. There appears to be no sufficient evidence to exclude
the possibility of a more brief incubation than ten days, the more
so when we consider that a formerly accepted minimum incubation
period of ten days for blood inoculation has given place to a later
minimum incubation period of two days.
III. AT WHAT TIME AND WHERE DID THE CASE
BECOME INFECTED?
It has been necessary to deal with a portion of this question
above in discussing incubation periods, but there are other facts
which may be employed to throw light on the subject.
Some assistance may be obtained from a study of the crescents
and sporulating forms. Thomson (1911) states that, as a general
rule, crescents appear in the peripheral blood on the fifth day after
the attack of fever.
Marchiafava and Bignami (1894) found crescents in the finger
blood of thirteen primary cases, usually between the seventh and
eighth day of the disease, rarely as soon as on the fifth. Bignami
and Bastianelli, who examined cases by spleen puncture, found
crescents in the spleen as early as the seventh and eighth day, and
exceptionally also in the peripheral blood.
It is seen from these observations that the spleen did not contain
crescents any earlier than the peripheral blood. This corresponds
with the findings in the present case, because an examination of
smears of peripheral blood, spleen and bone marrow showed that
whereas crescents were scanty in both peripheral blood and spleen,
they were very numerous in the bone marrow. From this fact
alone it could be deduced that death had occurred too early for
crescents to be liberated from the bone marrow in large numbers.
It has been shown by Marchiafava and Bignami (1894) that in
certain cases of fatal malignant malaria, crescents may be scanty in
the peripheral blood, spleen, and other viscera, while a very large
preponderance of them is found in the bone marrow, where they
may be seen in all stages. They conclude that the bone marrow
69
forms, if not the only, at least the most suitable site for the growth
of crescents. They compare the passage of crescents from the bone
marrow, where they are formed, with the corresponding passage of
nucleated red cells from the bone marrow where they also are
usually generated, into the general circulation.
Dudgeon and Clarke (1918-19) did not find crescents in their
cases; they state in general of their fifty-one cases that no histological
changes worthy of special reference were noted in the thyroid, bone
marrow, testicles and pituitary body. They cite a very acute case
of the haemorrhagic type, but no special mention of the condition
of the bone marrow is made; it is a significant fact, however, that
this case died in three days.
Applying the crescent periods and incubation periods to the
present case we have a sequence, tracing back from the day of
death, as follows:—October 12th, day of death, crescents present
in small numbers in blood; allowing five days for crescents to
appear in peripheral blood after paroxysm gives October 8th as
day on which in a normal case the paroxysm should have occurred;
allowing four days before this as the time when sporulating
parasites first appeared in the peripheral blood, brings us to
October 4th. From September 27th to October 4th is left for
parasitic incubation period on the theory that she became infected
by mosquito bite in the health resort in which it is known she was
exposed to anophelines. This parasitic incubation period (seven
days) is, even on the figures derived from the available experi¬
mental evidence, a probable one, and the history of the case supports
it. It should be recalled that in this patient no shivering or
sweating occurred and that we are dealing with a case of poor
physique and considered to have been anaemic, the effect of which
conditions upon incubation and reaction may be highly important,
although by no means sufficiently recognised. Incubation will
necessarily vary with the rapidity of the cycle of development of
the parasite, the special reproductive energy of the particular
parasite involved and the capacity of the infected human organism
to facilitate or impede such parasitic reproduction.
In the light of our present knowledge of incubation periods, the
occurrence of a mosquito infection six years ago or an inoculation
infection eight months ago seems improbable, and we have further
70
no evidence at present to warrant us in regarding the case as one
of endemic malaria. We are, therefore, compelled to decide
whether she.was infected at Liverpool or in the resort referred to.
The evidence appears to me to support the last named as the place
of infection.
THE SIGNIFICANCE OF CRESCENTS
Thomson (1911) considers the development of immunity to be
necessary before crescents can be produced, but he thinks that they
arise from ordinary merozoites, the period required for full develop¬
ment from merozoites to crescents being about ten days. He found,
in common with previous observers, that crescents appeared in the
blood about five days or more after the paroxysm, but while
accepting the existence of a condition of immunity as essential to
their production, he does not attribute their delayed appearance to
the lapse of time required before immunity is established but simply
to the length of time required for their development. The idea
that the delay was due to the time required for immunity to estab¬
lish itself had been put forward by Marchiafava and Bignami (1894)
partly and also by Stephens and Christophers (1908). From a
study of the appearances in this case, it is clear that the crescent-
producing process was more advanced in.the bone marrow than in
the spleen or blood, and if the crescent-producing process is an
‘ immunizing 9 one, we may safely assume that its effect was first
felt by the parasites in the bone marrow. Some evidence which
might support the view that the process is connected with
immunization may be obtained from a comparison of the relative
numbers of crescents and segmenting forms present in the peripheral
blood, spleen and bone marrow, respectively. In the peripheral
blood the number of crescents and segmenting forms was about
equal, in the spleen the segmenting forms far exceeded the crescents
(fifteen to one), while in the bone marrow the segmenting forms
were far outnumbered by the crescents (one to ten). Unless we
assume that segmentation occurs at different hours in the bone
marrow and spleen, or that red corpuscles containing young tropho¬
zoites cannot enter the bone marrow, on neither of which points
is there any evidence, we are compelled to conclude that some
agency in the bone marrow which causes the production of crescents
7 *
also prevents the completion of segmentation; that is to say, the
bone marrow is the seat of a process which hinders the asexual
while it facilitates the sexual development.
The number of crescents produced in any area such as the bone
marrow which contains only a proportion of the total blood must be
so small relatively to the areas where segmentation is occurring
freely, that it seems evident that a certain time must elapse before
the crescents reach the visibility point in the peripheral blood.
This, in my opinion, is the real cause of the delay in their
appearance, not only in primary cases but in relapses, and the delay
seems bound to occur, even supposing that the ‘ immunizing 1 process
already exists or becomes operative immediately in £he bone marrow
or other organs. It appears that this delay must occur, and that it
is not necessary to attribute it to a growth period of the parasite,
of the duration of which growth period this very delay is the only
evidence. In this case the 4 immunizing * process had commenced
in the bone marrow, but there was no evidence, as indicated by
great crescent production, that it had commenced in the other
tissues available for examination. It is possible that in other cases
it may occur early in other organs also.
NATURE OF THE ‘IMMUNIZING* PROCESS WHICH RESULTS
IN THE PRODUCTION OF CRESCENTS
Nothing definite can be deduced as to what this process is or
what are the factors which call it into operation. But that it is in
many cases quickly operative in the bone marrow is obvious, not
only from this case but also from the observations mentioned above
of the early appearance of crescents in bone marrow. It is also v
clear that many primary cases in which this process is already
established in the bone marrow, nevertheless die, which supports
the view that if the process is an ‘ immunizing * process it is focal
in origin.
That the process is initiated by the mere presence of, or rupture
of, segmenting forms, or by the toxins or pigment liberated at
sporulation, is unlikely, in view of the observation in this case that
in the spleen, where the most numerous segmenting forms occurred,
crescents were scanty.
72
SUMMARY
1. A patient who had never been out of the British Isles died
of malignant tertian malaria in Liverpool on October I2th, 1920.
2. There is evidence that this was an acute primary attack.
3. There is no evidence to show that this case acquired infection
by other means than mosquito bite.
4. From a consideration of the records of incubation period and
crescent formation, it is probable that the infection was acquired on
or about September 27th, on which occasion she was in a northern
health resort where • anophelines are plentiful (A. maculipennis ,
A. bifurcalus and A. plumbeus ). ♦
5. Some evidence is given that crescent formation commences
in the bone marrow, and that it is accompanied there by a failure
of complete development in the asexual forms.
6. The 1 immunizing ’ process which causes the above effects in
all probability commences in some cases very early in the infection,
but the crescents indicative of this process do not appear in the
peripheral blood for some time.
7. The late appearance of crescents in the peripheral blood in
infection with P. falciparum is explained on the ground that the
source from which they arise is limited in extent.
REFERENCES
Bastianelli, G. f and Bxgnami, A. (1899). Atti della See. per gli Studi d. Malaria , Vol. I, p. 28.
Craig, C. F. (1914)* Osier and McCrae's Modem Medicine, p. 73.
Dudgeon, L. S., and Clarke, C. (1918-9). Quart. Journ. Med., Vo). XII, pp. 372-389.
Glynn, E. G., and Matthews, J. C. (1920). British Medical Journal, pp. 811-813, Nov. 27.
Grassi, B. (1920). Quoted by Sella, Internal. Journ. Pub. Health, Vol. I, No. 3, p. 321.
James, S. P. (1920). Malaria at Home and Abroad, p. 150.
Jancs6, N. (1908). Atti della Soc. per gli Studi d. Malaria, Vol. IX, p. 143.
Marchiatava, E., and Bignami, A. (1894). Syd. Soc. Monog. on Malaria, pp. 64 and 211.
Mitemain, M. B. (1916). U.S.A. Pub. Health Rep., Vol. XXXI, No. 6, p. 301.
Osler, W. (1901). Clifford Allbutt’s System of Medicine, Vol. II, p. 730.
Ross, R. (1920). British Medical Journal , Dec. 4, p. 871.
SchOitnir, W. (1901). Zeitscbr.f. Hyg. u. Injektiontkr, Vol. XLI, pp. 89-122.
Stephens, J. W. W., and Christophers, S. R. (1908). The Practical Study of Malaria and
other Blood Parasites, p. 52.
- and Others ( 1917). Ann. *Trop. Med. and Parasit., Vol. XI, pp. 103-110 and 158-164.
Thayer, W. S., and Hewktson, I. (1895). John Hopkins Hospital Rep., Vol. V, p. 35.
Thomson, D. (1911). Ann . Trop. Med. and Parasit ., Vol. V., p. 57.
73
OBSERVATIONS ON MOSQUITOES IN
THE ISLE OF MAN
BY
B. BLACKLOCK
AND
H. F. CARTER
(Received for publication 22 February , 1921)
Plates X—XIV
In recent years much interest has been shown in the mosquitoes
of the British Isles, and many new observations have been made
which increase our knowledge of the species which occur here.
Twenty-two species, including three members of the genus
Anopheles , have now been recorded from the United Kingdom, and
many of them are also known to occur in Ireland; but, so far
as we are aware, no information concerning the mosquitoes of the
Isle of Man is available. Having recently had occasion to visit the
Island, we took the opportunity of investigating the occurrence and
distribution of mosquitoes. As, however, our visit extended over
only a portion of November and December, it was impossible to
make any detailed investigation of even those mosquitoes of which
the over-wintering stages are known and readily found. Our
activities were for the most part confined to localities easily
accessible by railway or tramway, and were, therefore, mainly
limited to the coastal region. The places visited and the records
obtained are shown in the appendix and map.
GENERAL CONSIDERATIONS
I. Anopheline Mosquitoes.
A. bifurcatus , L. The larvae of this species were abundant and
widely distributed over the Island, and were found without difficulty
in spite of the fact that residual breeding-places only could be
discovered at this time of the year. As is sometimes the case with
mosquitoes of this kind, larvae were found in certain of the
breeding-places only after considerable searching; this was especially
noticeable in the more extensive breeding-places, such as the marsh-
74
land (Photo. No. ij in the neighbourhood of Ballaugh and Sulby.
On the other hand, relatively large numbers were sometimes obtained
in a short time in the more circumscribed waters, such as the
artificial pond at Groudle Glen (Photo. No. 2) and in the ditch at
Castletown (Photo. No. 3). In the lattfcr case, in the small ditch
shown on the right of the photograph, over sixty larvae were
captured in ten minutes by one person using a small scoop. As will
be seen in the photographs, breeding-places of A. bifurcatus were
found in various types of country, even close to the sea (Photo.
No. 4); they occurred also in situations where free water was only
discovered by close inspection and was chiefly limited to hoof
prints, etc. (Photo. No. 5). They were also found in the near
neighbourhood of towns, as shown in Photo. No. 6 (Appendix,
No. 15).
A. maculipennis , Mg. Hibernating females of this mosquito
were not found in such large numbers as (in view of the numerous
potential breeding-places observed) we expected. All types of
cow-sheds, stables, and other out-buildings from those of
modem construction—high-roofed, well-lighted and well-ventilated
—to those of primitive form—small, dark and badly ventilated—
were systematically examined; it was naturally impossible, however,
to obtain access to houses, and, therefore, it is not -possible to give
records relating to them. In only one case (Appendix, No. 21,
Photo. No. 7) was any considerable number of females found, when
one hundred and forty-seven were captured, this representing the
great majority of those present in the building. In accounting for
the relatively small numbers seen, the following two facts, which
came under cur notice, are possibly of some importance: —
1. Cleaning and lime-washing. In several instances we were
informed that a thorough sweeping of the walls and roof to remove
cobwebs had recently been carried out; this procedure in some cases
had been followed by lime-washing.
2. Accidental cleaning of the roofs. Having observed in some
cowsheds and stables with low flat roofs that cobwebs existed only
in the comers, or over the manger, we made enquiries as to the
reason of this. It is to be attributed to the practice of carrying on
a fork the hay used as fodder, thus repeatedly brushing the greater
portion of the roof.
Annals 'Trap. Med. & Parasitol .. Vol. AT
PLATE X
Photograph No. i
Photograph No. 2
C. Tntling & Co., Lid., Imp
75
A. plumbeus , Steph. Although the Isle of Man is by no means
heavily wooded, we experienced little difficulty in discovering the
larvae of A. plumbeus on numerous occasions. The breeding-places
were similar in nature to those of which we have given illustrations
in our previous papers (1920). We found here a surprising
difference in the tendency of the Spanish chestnut ( Castanea
saliva) to form rot-holes, and although no systematic survey was
carried out, yet of nineteen breeding-places discovered, four
occurred in this species of tree; whereas in Liverpool and district,
of sixteen breeding-places found, none were present in the Spanish
chestnut. Photo. No. 8 displays the most interesting breeding-
place of this mosquito that we have yet discovered; a full description
of it is given on p. 87 (No. 34). Interesting breeding-places were
also found immediately adjacent to an out-door latrine (Photo.
No. 9) at a country farmstead, and near the centre of a town as in
Photo No. 10 (Appendix, No. 31 f).
It is necessary to note here that in no instance were the larvae of
0 chlerotatus geniculatus , Ol., discovered. So far as our investiga¬
tions have gone, although we have commonly found this species in
the United Kingdom in association with A. plumbeus , we have
entirely failed to find it either in Ireland or the Isle of Man,
although A. plumbeus is abundant in both places.
II. Culicine Mosquitoes.
Culex piptens , L. Females of this species were found in several
of the localities in which buildings were visited. They were
frequently found associated with A. maculipennis in occupied cow¬
sheds, stables and piggeries, but were distinctly more numerous in
cooler stituations, i.e. t cellars, lofts, out-houses, etc. (Appendix,
p. 88).
Theobaldia atinulata , Schr. Larvae, pupae and adults of this
mosquito were discovered in various places. The adults were
relatively uncommon, but females were occasionally found in
buildings, and, on one occasion (November i8tE), both sexes were
emerging from pupae contained in a wooden tub. This was
standing in the back garden of a cottage at the Nunnery Mill,
Douglas; the position of the tub in relation to the cottage is seen
in Photo No. 11. The dimensions of the tub were 18 inches in top
76
diameter and 12 inches in depth; it was nearly full of rain water,
which, on being stirred, produced a very evil odour due to a mass
of putrifying leaves lying on the bottom; no larvae were found.
Eighty-four pupae were collected from the tub, and from seventy-
eight of these, twenty-seven males and fifty-one females emerged.
Pupae of T. annulata were also found at Ramsey on November 22nd,
in a marshy meadow situated behind the town (Photo. No. 6).
Larvae of the second, third and fourth instars were abundant, and
occurred in several different types of breeding-place (Appendix,
p. 88 ).
Culicella morsitans , Theo. Larvae of various sizes were found;
they sometimes occurred with the larvae of T. annulata , but were
more definitely limited in regard to habitat. They were always
found in open water containing abundant vegetation, and, in
marshy land, appeared to favour more particularly the immediate
neighbourhood of clumps of rushes (Appendix, p. 89).
Culicella fumipennis , Steph. Larvae of this species were found
in one locality only, namely, in the extensive peat bog-land or
' curragh * in the vicinity of Ballaugh. They were, however, taken
in two sites, but the nature of the breeding-places in regard to the
types of vegetation present and colour of the water (reddish-brown)
appeared identical. These larvae, some of which appeared but
half-grown, were of a characteristic yellowish colour, and could
immediately be distinguished from those of C . morsitans , which
were darker and distinctly brown (Appendix, p. 89).
APPENDIX, GIVING RECORDS OF SPECIES OBTAINED, UNDER
PLACE NAMES ARRANGED IN ALPHABETICAL ORDER
Anopheles bifurcatus , L.
( 1 ) Ballaugh, December 6th.
(a) The country immediately north of the railway to the East
of Ballaugh Station presents features which are peculiar to this
portion of the Island. It is a flat, desolate marshy country, with
large expanses of water interrupted by collections of low bush and
reeds. In Photo. 1, which gives a good idea of the appearance of
much of this region, known as * the Curragh,* the foreground repre¬
sents an area adjoining the path, completely waterlogged and
Jr rails Trop. Mod. £> Paras itolVo!. XV
PLATE XI
Photograph No. 5
C. Tinting c 1 Co., Ltd., Imp.
77
communicating with the open water beyond. Larvae were found
among the coarse grass seen in the foreground of the picture, but it
was impossible to make any observations on the open waters.
Besides this breeding-place, two others were found of quite a
different character.
( b ) One was a pond in a field where the peat formation of the
district was observed at the edges of the pond itself, and also in
the brown discolouration of the water. Here larvae were obtained
after breaking ice one-eighth inch thick.
(c) The other was near the railway line close to the station, and
was a cutting for water, having a high bank, formed by the railway
track, on one side. The water was clear and flowing, but in shallow
areas at the margin, where grass, duckweed and watercress provided
shelter, larvae were numerous.
(2) Castletown, November 30th.
(a) Near the public park, and separated from it by the river
(Silver Bum), is a meadow with a stream running through it; this
stream at the time of our visit was overflowing at various points,
covering a large area with shallow, stagnant water. Only at one
end of the flooded area, in a boggy patch, with fioofprints, was a
single larva obtained after considerable searching, and none was
found in the stream itself, which was moving fairly rapidly.
(b) In striking contrast to the absence of larvae from the centre
of the meadow, was their abundance in a narrow ditch on the side
remote from the river (Photo. 3).
In the photograph the ditch in question is on the right, while the
flooded area extends to the left for a considerable distance outside
the picture. In this ditch, which varied in width from 3 feet to
about a foot in some parts, and which was a foot deep with over¬
hanging banks, sixty larvae were collected in ten minutes by one
person using a small scoop. The vegetation was chiefly grass and
watercress, and from the abundance of life of various kinds present
we were led to conclude that this formed a permanent and very
suitable breeding-place for A. bifurca/us.
(3) Douglas, November 18th.
In the neighbourhood of Douglas outside the Nunnery estate,
about twenty minutes' walk from the quay, an extensive breeding-
78
ground is present. It is that portion of flat land which lies to the
right of the path outside the Nunnery towards the racecourse. The
only indication of the marshy nature of this ground was the
appearance of reeds, as the water was concealed by the vegetation
and was not visible even at a few yards distance.
The area is approximately 120 yards by 35 yards, and is
continuous with a similar area not examined by us on the left of the
path, where it turns towards the racecourse. In the area under
consideration larvae were constantly found on testing in various
directions at five-yard intervals, very frequently in hoofmarks.
Beyond this field, across the hedge towards the racecourse, is what
appeared at first sight to be a continuation of the same sort of half
marshy ground, but the water present was red and contained a great
quantity of flocculent material. No larvae were found here despite
prolonged testing.
(4) Glen Garwick Road, November 19th.
No larvae were found actually in this glen, although a swampy
patch just in front of the hotel was regarded as promising, but a
breeding-place occurs beside the Electric Tramway line from
Groudle, just beyond the level crossing past Liverpool Arms Hotel
to the left of the line going towards Garwick. Larvae were found
here in a field which was marshy, and their distribution appeared
to coincide with the occurrence of patches of duckweed.
(5) Glen Greenaugh, December 1st.
In a pond on the property of Balia Vale, just above the weir
across the stream, larvae were found in small numbers. The pond
contained a large quantity of dead leaves, and at one end where
larvae were found grass was present; it is situated about 150 yards
from the house.
(6) GLEN Groudle, October 28th and November 16th.
(a) On the occasion of the preliminary visit, a large breeding-
place was found in the glen in the shape of an artificial pond which
lies between the path and the river, and which is used as a fernery.
The breeding-place is 80 yards long by 25 yards across at its wfdest
part, and is situated about 500 yards from the viaduct. The pond
is sub-divided into what is practically a series of small ponds by
Annals Trop . Med. & Parasitol., VoL XV
PLATE XII
Photograph No. 6
Photograph No. 7
C. Tinting <5^ C 0 ., Ltd.. Imp
79
means of banks of considerable width. Larvae were found with
ease all over the area. The vegetation consists of coarse grass,
reeds, waterlilies and, on the banks, ferns (Photo. 2).
(J>) On the second visit, by passing down the glen, we found a
breeding-place at Groudle Creek in close proximity to the sea,
separated from it, in fact, only by a bank of shingle and the banks
of the river itself; it is seen in Photo. No. 4.
From its proximity to the sea and from the presence of much
seaweed and flotsam in this collection of water, we concluded that
it must contain a considerable excess of sodium chloride, and the
water had a brackish flavour; but analysis* showed that, in fact, it
contained only 7*63 parts per 100,000, which is stated to be lower
than many potable waters of the Isle of Man.
This breeding-place, which was about 30 yards from the sea and
100 yards from the nearest croft, did not yield numerous larvae.
(7) Glen Laxey, November 20th.
Close to the entrance to Laxey Glen Gardens there is a boggy-
looking field on the left of the path, in which larvae were found.
The field is bordered by a wall (seen in Photo. 5), which separates
it from the bandstand and pleasure ground. To the right of the
position seen in this photograph are various swings and apparatus
for the amusement of the numerous children visitors. The hotel lies
behind the camera to the right at a distance of about a hundred
yards.
It may be noted again here that even at a short distance the
presence of water could not be perceived, but could only be inferred
from the character of the vegetation.
(8) Glen Sulby, December 7th.
The Curragh region mentioned above in connection with Ballaugh
extends towards Sulby Glen, and can be approached from this
point.
0 a ) At the edge of the Curragh larvae were found in a field
surrounded by stagnant ditches. The centre of the field was
swampy and larvae were found here, and, where the water was clear
among the reeds, in one of the ditches bordering the field; in two
• Kindly carried out for u« by Mr. Fyffe of the Public Analyst* Laboratory, Douglas,
Isle of Man.
8o
other ditches, where the water was loaded with red flocculent
material, no larvae were discovered.
( b ) Another breeding-place was found in Sulby itself, about
400 yards from the Glen Hotel. It was a marshy piece of ground
beside the road, and was bordered on the far side by an area of
higher ground where many trees had recently been cut down. The
photograph showing this breeding-place of A. bifurcatus shows also
a very uncommon breeding-place of A. plumbeus , and will be found
under the latter heading, Photo. 8.
(9) Glen Willyn, December 7th.
In the terminal part of this glen, about 300 yards from the sea,
there are pleasure grounds furnished with bandstand, swings, etc.
These abut on the small stream, and just opposite them is a narrow
wet strip of ground at the foot of a steep hill. In this wet area
larvae were found in that portion adjacent to the fence which runs
along the side of the stream, and separated from the pleasure ground
by only the width of the stream.
(10) GREEBA, December 5th.
The railway from Douglas to Peel rims in a valley for a consider¬
able portion of the way. The land close to the railway is marshy,
and has at intervals much free water lying on it. This strip of wet
land was examined in the region of Greeba, and there was no
difficulty in finding larvae.
(11) Peel, December 8th.
Larvae were found in a small narrow drain at the foot of the hill
bordering the road along the quay on the way to Peel Castle. The
water was sluggish and much grass was present in many places.
(12) PORT Erin, November 26th.
Three breeding-places were discovered in this locality.
(a) The first was in marshy ground in the valley near the far end
of the golf course.
(b) The second was a slow-moving stream with high banks and
much vegetation, chiefly grass and watercress situated between Port
Erin and Ballafesson village.
(f) The third was a small piece of marsh land covered with reeds
in the centre of the village of Ballafesson.
Annals Trop. Med, & Parasitol .. Vol. XV
PLATE Kill
Photograph No. 9
Photograph No. S
C. Tinling &* Co., Ltd., Imp.
11
(13) Port Soderick, December 5th.
About half a mile before Port Soderick, on the road from
Douglas, beside the road a breeding-place was found. It consisted
of a pool of clear water, deep in the centre with a rapid streamlet
running through it. The vegetation was considerable, and consisted
of a fairly thick layer of duckweed which covered the centre of the
pool and reached to within a few inches of the edge in most places.
The edges were covered with grass and duckweed. The larvae were
present in numbers, and were found chiefly near the edges.
(,14) Port St. Mary, November 29th.
Close to the station, in a meadow belonging to Ballacreggan
Farm, just behind the gas works, a small ditch a foot wide, over¬
grown with grass and not conspicuous, proved to contain larvae.
The water was clear and had a few patches of duckweed on it at
intervals, and the surface of the water was about a foot below the
level of the banks, which were undermined.
(15) Ramsey, November 22nd.
An excellent breeding-place was encountered on the outskirts of
the town. The extent of this area, a marshy field, is 120 by
120 yards, approximately. Very numerous larvae were found here.
Situated about 15 yards away is the playground of a large school,
which can be seen in Photo. No. 6 . In the whole of this area larvae
were found.
(16) ST. JOHN’S, December 8th and 9th.
(а) Larvae were found in a pool in a field adjacent to the
station. This pool serves partly as a cesspool for drainage from
the station lavatories; it is situated 10 yards from the platform,
6 yards from the lavatories and about 30 yards from the main road.
The water was slightly coloured, the margins very boggy, and the
vegetation mostly rushes with some duckweed and grass.
( б ) On the road from St. John’s to Peel, in a field on the right
Side of the road about half a mile out of St. John’s, larvae were
found in the margins of a narrow, rapid stream at the foot of a hill.
(c) Further on the road, about a quarter of a mile beyond this
breeding-place, a considerable amount of marshy ground exists at
82
the side of the railway. There is an extensive breeding-place
between the railway and the St. John’s-Knockaloe Road, the edge
of the marshy ground lying ioo.yards distant from the nearest
farmstead.
Anopheles maculipennis , Mg.
A number of cowsheds, stables, piggeries, fowl-houses, latrines,
outhouses, cellars, lofts, sheds and shelters of various kinds were
examined with a view to finding the adults of this mosquito.
(17) BALLAUGH, December 6th.
(a) Three cowsheds were examined, of which two were
small, dark, warm, flat-roofed, with numerous cobwebs
present, and the third lofty, well-lighted, with gable roof
and almost free from cobwebs. In one of the former, three,
and in the last, one A. maculipennis 9 were captured. Total 4 9
( b ) One stable of modem construction, with high gable
roof, but rather dark . 29
(18) Castletown, November 30th.
(a) One cowshed, small, dark, warm, low-roofed, situated
in the centre of the town . ... 10 9
( b ) One stable for two horses, in centre of town, low-
roofed and warm, with few cobwebs except in one comer ... 39
(1 c ) One outhouse adjoining stable, and of same construc¬
tion, used as a potato store, cooler and draughty . nil.
(d) Two lofts; one warm, dark, slate-roofed, with few
cobwebs, situated over (b) and ( c ), nil; the other situated
above ( a ), dark, slate roof, numerous cobwebs, I 9 • Total 1 9
(19) DOUGLAS, November 18th.
At the Nunnery Mill the following were examined: —
( a ) Stable, a large, dark stable on ground floor, just being
whitewashed, few cobwebs . nil.
(b ) Fowl-house, low and dark ... . nil.
(r) Cellars extensive, low-roofed, under mill house ... nil.
( d ) Loft situated above (< a ), but not newly whitewashed,
cobwebs numerous . nil.
( e ) One stone-built shelter, used as a fowl run . nil.
Annals Ttop. Mai. if Parasite!.. Vol. XV
PLATE XIV
Photograph No. io
Photograph No. i i
C. Tinling Co., Ltd., Jrr.p.
83
At the back of the town: —
(a) Stable, small, dark, flat-roofed, warm, numerous large
cobwebs . . nil.
(b) Outhouse adjoining, lighter and cooler . nil.
(c) Loft above (a) and (b), dark, warm, wood-roofed ... nil.
(20) Glen Garwick Road, November 19th.
(a) One stable, disused . .. nil.
(£) One loft, above (a) . . nil.
(21) Glen GROUDLE, October 30th, November 15th and 16th.
(a) One cowshed, situated 15 yards from a cottage
(Photo. 7) and 100 yards from a breeding-place of
A. bifurcatus was examined. It was gable-roofed but dark,
low, warm and had many cobwebs. The door was central in
the side wall, and at one end two cows were kept, the other
end being a coal and wood store. There were captured here,
distributed equally over both ends of the shed, but less
numerous just at the door .147 $
( b ) One stable for two horses, dark, flat-roofed,
moderately warm, but draughty.22 9
(c) Three latrines near stable ... ... ... ... nil.
id) Four cellars situated near stable, two in house and
two under verandah . nil.
( e ) Three outhouses .'.. nil.
(/) One loft over ( b ), communicating with the stable by an
opening two feet square above the manger, and by ladder
entrance also, well-lighted, clean and airy, recently renovated
roof of corrugated iron, few cobwebs, draughty ... ... nil.
(g) Two sheds, corrugated iron, a few yards from stable,
containing piles of wood. nil.
(h ) One shelter (public) in glen . nil.
(i) One kennel . nil.
(22) Glen Sulby, December 7th.
Two cowsheds: —
(a) One very small, two animals, gabled, corrugated iron
roof, dark, warm, the roof moist, cobwebs very numerous ... 99
(b) Similar but wood-roofed, recently swept, few cobwebs nil.
8 +
(23) Kirkmichael, December 7th.
(a) Three cowsheds, flat-roofed wood, dark, warm, clean,
to hold twenty, eight, four animals, respectively . nil.
( [b ) One pigstye, usual type, dark, fairly warm, cobwebs
numerous.. . nil.
(c) One loft above largest cowshed, high gable roof,
dark, warm, many cobwebs . nil.
(24) Peel to St. John's, December 8th.
(a) Two cowsheds: —
(i) Large, sixteen animals, flat-roofed, warm, cob¬
webs over mangers and in corners only ... ... 49
(ii) Large, ten animals, flat-roofed, dark, few cob¬
webs, except in comers ... ... . ... 17 9
( b ) Two stables: —
(i) Flat wood roof, warm, dark, few cobwebs ... 29
(ii) Similar but more cobwebs . nil.
(1 c ) One latrine outside house ... ... ... ... nil.
( d ) One loft above (a) (ii); high gable roof, cool and airy,
containing straw and hay . nil.
(25) PORT Erin, November 26th and 29th.
( a ) Four cowsheds: —
(i) Two, low flat wood roof, dark, warm. nil.
(ii) Two, iron roof, low, warm, moist, dark ... nil.
( b) Three stables, dark, unventilated, cooler than cow¬
sheds. In one . 1 9
(c) One pigsty, low, dark, wood roof, many cobwebs ... nil.
(d) Two fowl-houses, thatched low ... . nil.
(e) One loft above (a) (i), dark, cool ... ... ... nil.
(26) Port St. Mary, November 29th.
(1 a ) Two cowsheds, large, dark, gable, slate roof recently
limewashed, few cobwebs . nil.
(£) One outhouse, continuation of cowshed, cold, draughty nil.
( c ) Two sheds, one for straw, one for carts . nil.
(27) RAMSEY, November 22nd, 23rd and 24th.
(a) Four cowsheds:—
(i) One for six animals, low wood roof, dark,
warm . . 19
(ii) Three, first rather light, cool, corrugated iron
roof; second and third warm, low wood roof, dark;
in third . 39
( b ) Four stables: —
(i) One large, airy, high corrugated iron roof, few
cobwebs, accommodation for about fifty horses ... 1 9
(ii) Two, small, wood roof, dark, with cobwebs... nil.
(iii) One, airy, light, cool, wood roof, cobwebs
numerous ... . ... . nil.
(0 Six pigsties; usual type, in each of two, one female.
Total ... 29
Anopheles plutnbeus, Steph.
(29) Ballasalla, December 9th.
Sycamore. A breeding-place was found just outside the
churchyard of the Abbey Church, between it and'the river. The
tree was situated a few yards from the road. The aperture was at
a height of 5 feet from the ground and i| inches in diameter, depth
of hole 6 inches; the water was yellow tinged.
(30) Ballaugh, December 6th.
Sycamore. In the farmstead of Mr. Kneen a breeding-place of
this mosquito was found in a tree situated about 30 yards from the
dwelling-house. The aperture, situated 8 feet from the ground,
was 6 inches in diameter and the depth of the cavity 12 inches.
Small larvae only were found.
(31) Douglas, October 30th, November 18th, December 3rd.
Seven breeding-places were found in and near Douglas. On the
previous visit, October 30th, two breeding-places, (a) (b), were
found in the small glen behind Falcon Cliff Hotel.
October 30th: —
(a) Sycamore, situated 15 yards from the garage and 27 yards
from the main road. Aperture at 1} feet from the ground, diameter
86
about 6 inches, depth 9 inches. The whole trunk was hollow.
Large and small larvae numerous.
(J?) Sycamore , 60 yards from garage and 10 and 12 yards,
respectively, from two roads. Aperture 1 foot from ground, hole
9 inches deep; small larvae.
November 18th: —
(c) Sycamore in garden of Nunnery Mill Cottage, situated
12 yards from house. Aperture 12 feet from ground, measuring
2 by i| inches, numerous larvae.
( d ) Lime in paddock below cottage and mill. Situated 30 yards
from house and 15 yards from the mill.
( e ) Sycamore just outside garden of Mill Cottage. Aperture
about 4 feet from ground, with a smaller aperture 3 feet from the
ground almost level jvith the water. Owing to difficulties in either
siphoning or scooping out the water in this cavity, resort was had
to flushing out the larvae by pouring a bucket of water into the
upper opening and catching the water as it emerged from the lower
opening.
December 3rd: —
(/) Sycamore situated in garden of Marina Terrace, at the
junction of two streets and distant 20 yards from an elementary
school (Photo. No. 10). Aperture at 2 feet from ground 5 by
3 inches in size and cavity 6 inches deep.
(g) Sycamore in same garden 5 yards from the previous one.
Aperture 2 feet from ground 8 by 5 inches, depth 2 feet; small
larvae numerous.
(32) Glen Groudle, November 19th.
Spanish Chestnut . Just where the bridge on the high road
crosses the Groudle river, a very large breeding-place was found.
The aperture, which is at 10 feet from the ground, appears over
the parapet of the bridge and is 9 inches in diameter, the cavity
being full of red water to within 2 inches of the lip of the aperture.
(33) Laxey, November 20th.
Spanish Chestnut . In Laxey Glen Gardens, on the right side
of the arena in front of the bandstand, are some trees on a slightly
raised terrace; in one of these, larvae were found; aperture at feet
87
from the ground, 3 inches in diameter and facing upwards, depth
of cavity 15 inches.
(34) Glen Sulby, December 7th.
Ash . Breeding-place found in this locality is of exceptional
interest. In an area situated 400 yards from the Glen Hotel, where
many trees had recently been cut down, larvae were found in the
stump of an ash tree. The aperture is a foot in diameter, but before
water is reached at 18 inches from the top of the stump the c'avity
has widened jjreatly, so that the surface -of the water is very much
larger than the aperture of entrance. Not only so, but also the
cavity extends downwards until it is much below the level of the
ground. That the breeding-place is a true rot-hole, and entirely
contained in the tree stump, is shown by the facts that it was
impossible to push a stick down through it at any place, and that
the water was deep red in colour; it contained numerous larvae of
Myiatropa florea and various chironomids (Photo. No. 8). This
photograph is further of interest because the marshy ground below,
indicated by arrow, is a breeding-place of A. bifur cat us.
(35) Kirk Braddan, December 10th.
Sycamore. Near Braddan Church, a tree was found with a
cavity containing larvae. It stood on a bank behind the Hotel at
a distance of 20 yards. Two apertures were present about 4 feet
from the ground and about 5 inches in diameter.
(36) Kirk Michael, December 7th.
Spanish Chestnut. The breeding-place found here was in a tree
situated in a field to the right of the road leading to a farm
(Mr. Mylchrcest), and distant 12 yards from the cowsheds and
80 yards from the White House. The aperture was at a height of
7 feet and was remarkably small, about 1 inch. By means of the
siphon, however, a great rush of red water was obtained from a
large cavity, with larvae in numbers.
(37) Peel, December 8th.
Spanish Chestnut. The nearest point at which A. plumbeus was
found was the estate of Ballamore. The tree was situated beside
the main road. The aperture was 9 feet from the ground and
8 inches in diameter, depth 6 inches.
m
88
(38) RAMSEY, November 23rd.
On the estate of Miltown, outside Ramsey, two breeding-places
were found.
(a) Sycamore , situated 15 yards to left side of Ramsey Road.
Aperture at 10 feet from ground 6 by 7 inches, depth 2 feet, the
water surface being about 1 foot from lip of aperture; numerous
larvae
(J?) Sycamore. In close proximity to the foregoing, aperture at
8 feet from ground, 12 inches in diameter, a very large cavity full
of a pulpy mass of debris, with a layer of water on the top.
(39) St. John’s, December 8th.
At a house on the St. John-Knockaloe Road, just beyond
Ballamore estate on the left of the road, two breeding-places in
sycamores were found.
( a ) Sycamore . This was situated 20 yards from cowsheds and
30 yards from house; aperture at 4 feet from ground, 9 by 4 inches,
depth 9 inches; trunk above hole also hollow for more than 18 inches
up; numerous larvae, over twenty in the first scoopful of water taken
out.
(b) Sycamore . Standing immediately beside a latrine, 15 yards
from the house. There was a large cavity with two apertures, each
about 5 to 6 inches in diameter (Photo. No. 9).
Culex pipiens , L.
Hibernating females were found in the following places: —
Douglas, in 1 stable, 1 fowl-house, 1 cellar, 1 loft, 2 shelters,
1 latrine; Glen Garwick Road, 1 stable, 1 loft; Groudle Glen,
1 outhouse, 2 latrines, 2 cellars, 1 loft, 2 sheds; Port St. Mary,
1 shed; Ramsey, 3 cowsheds, 2 stables, 3 piggeries, 1 loft, 1 shed.
Theobaldia annulata , Schr.
Hibernating females were found: —Groudel Glen, 1 cellar, 1 loft,
1 latrine, 1 open shelter; Kirkmichael, 1 cowshed.
Pupae. Douglas, Nunnery Mill Cottage, in wooden tub (see
photo, n). Ramsey, in association with larvae of A. bifurcatus
(Appendix, No. 15).
89
9 °
Larvae. BaUaugh, under J-inch ice in peat bog and pond in
field; Castletown, ditch; Glen Garwick, concrete artificial pond;
St. John’s, manure pit; Port Erin, marshy field; Ramsey, see above,
under pupae.
CuliceUa morsilans, Theo.
Larvae. Ballaugh, under £-inch ice in pond in field and peat
bog, in association with T. annulata and C . fumipennis larvae;
Douglas, marshy field; Glen Garwick Road, marshy field; Glen
Sulby, marsh.
CuliceUa fumipennis , Steph.
Larvae. Ballaugh, as above, under C . morsitans.
REFERENCES
Blacklock, B., and Cartxk. H. F. (1920). Observations on Anopheles ( CoAodiazesis)
plumbeus , Stephens, with special reference to its breeding-placet, occurrence in the
Liverpool district and possible connection with the spread of malaria. Ann . Trop.
Med. and Parasite Vol. XIII, pp. 421-452.
Blacklock, B. and Castii, H. F. (1920). On the results ’obtained from surveys for
breeding-places of tree-hole mosquitoes in Liverpool and neighbourhood. Ibid., Vol.
XIV, pp. 115-126
9 1
NOTES ON SOME UNUSUAL BREEDING-
PLACES OF STEGOMTIA FA SC I AT A,
Fabr., IN AUSTRALIA
BY
G. F. HILL, F.E.S.
From the Australian Institute of Tropical Medicine ,
Townsville , Queensland
(Received for publication i March , 1921)
Plate XV
Although generally regarded as a domestic species, there are
several records of S. fasciata breeding in rot-holes in trees and in
water-retaining plants; this habit, however, appears to have escaped
notice hitherto in Australia.
On 20th January, numerous larvae and pupae of this species and
of Ochlerotatus notoscriptus , Skuse, were collected from a tin
containing about 5 inches of water and a quantity of decaying
leaves, which was found in the dense scrub 600 yards distant from
the nearest of several seaside dwellings on Magnetic Island. Several
adult Stegomyia which were captured at the same time (1 p.m.)
while attempting to bite, and others which were bred from the
larvae and pupae, were noticed to be distinctly darker and smaller
than the forms found throughout the year in the Institute buildings.
In order to determine the permanency of this dark form, a number
of these males and females were bred from for five generations under
the usual laboratory conditions, and concurrently with an equal
number of generations from individuals of the lighter coloured form
commonly found in dwellings. From both series males and females
of each generation were secured for comparison, the experiment
being terminated, in the case of the dark form, with the sixth
generation, which comprised males only. An examination of the
material thus obtained showed that the light form bred true,
i.e. t the individuals of each generation did not differ from their
progenators, whilst the dark form produced, in each generation
92
after the first, a proportion of individuals of both forms, as well as
intermediate forms. In both series some variation was noticed in
the shape and width of the outer lyre-shaped thoracic ornamentation
and in the length and width of the median thoracic stripes, but in
none were the latter entirely absent, a character recorded by Taylor
( 1914 ).
The island referred to above lies in Cleveland Bay, about four
miles distant from Townsville, and was formerly utilised as a site
for the port Quarantine Station.
On 5th May, a considerable number of mosquito larvae were
siphoned from a rot-hole in a poinciana tree (Plate XV), growing in
the Hospital grounds and distant about 70 yards from the nearest
dwelling. In addition to Macleava tremula , Theobald, and
Ochlerotatus quasirubrithorax, Theobald,* this collection produced
a large number of Stegomyia fasciata , similar in size and coloura¬
tion to the Magnetic Island form. On 7th September, another
batch of larvae and pupae were collected from this hole, and from
them about one hundred adults were bred. In this case the males
were of about the average size, but quite as dark as any I have
seen, whilst the females were of the light form and unusually large.
REFERENCE
Taylor, F. H- (1914)* Proceeding* of the Linnean Society of New South Wales. Vol.
XXXIX, p. +55.
Identified by Mr. F. W. Edwards, as a variety of this species.
-*?finals T:op. Med . of Parasitol / 7 o/. A’/
PLATE XV
ROT-HOLE IN POINCIANA TREE.
BREEDING PLACE OF OCHLEROTATUS QUASIRUBRITHORAX,
M ACI.EATA T RF.MULA AND STEGOMVIA F ASCI AT A
C. Tinling o Co., 7m/>.
93
MUSCA DOMESTICA, Linn., AS A
‘BUSH FLY’ IN AUSTRALIA
BY
G. F. HILL, F.E.S.
From the Australian Institute of m Tropical Medicine ,
Townsville , Queensland
( Received jor publication I March , 1921)
In the literature dealing with the etiology of M. domestica ,
I can find no record of this species in the r 61 e of a ‘ bush fly/ i.e., a
fly which lives and breeds normally beyond the range of human
habitations, although I understand that Major E. E. Austen is of
opinion that this species originated in the tropics and has thence
spread to temperate climates, where it is only able to maintain itself
by the fact that it has taken to living in houses.
In two widely separated localities in North Australia, evidence
has been gathered which strongly suggests in one case, and
proves conclusively in the other, that this ubiquitous species is
not dependent upon human habitations and environments for its
existence.
During the period 1913-1917, specimens were frequently
captured in the Darwin District (Northern Territory) on stock and
carcasses at distances up to a couple of miles from dwellings, but
for some time it was considered probable that these flies were bred
under the usual conditions; later, however, examples were captured
in company with Muse a ventrosa f Wied. ( M . nigrithorax , Stein.),
and M. humilis , Wied. ( M . vetustissima> Walk.) on the carcasses
of freshly skinned buffaloes which had been shot in scrub country
from three to six miles distant from the nearest habitation—a cattle
station sixteen miles distant from the next nearest dwelling and
about thirty miles from Darwin. During the summer months,
M. domestica were very numerous at the station, especially in the
kitchen and adjacent living room, but they were not seen on men
or horses after leaving the homestead for the haunts of the buffaloes.
In April, 1919, an officer of the Stock Department, Townsville
(N. Q.), brought in for identification a large number of flies which
had been captured in the vicinity of a slaughter-yard and upon
94
stock grazing in the locality, in some cases more than a mile from
the nearest dwelling. This and later collections invariably included
a large proportion of M. domestica .
During the months of April to October, flies of the same species
were frequently captured upon grazing horses and cattle, and upon
my face, hands and clothing in the Town Common at distances up
to two and a half miles from habitations. On these occasions they
were generally associated with M . lusoria , Wied. 1 (M. australis ,
Macq., 2 M. fergusoni , J. & B. 3 ), and M. nebulo , F. 4 , ( M . hilli ,
J. & B. 6 ), the last named being less aggressive than the others.
From October to about the end of March, M. humilis , Wied. 6
(M . vetustissima , Walk.) is the predominant species, and is
certainly the most widely distributed Musca found in Australia,
being as plentiful in the outer suburbs of Melbourne (Victoria) as it
is in Central Australia, N. Territory, and the N.W. and Kimberley
Divisions of W. Australia.
In the bush or open grazing country near Townsville,
M. domestica oviposits on fresh horse-droppings, but they will also
oviposit and rear their progeny on decaying vegetable matter, as
shown by the fact that upon two occasions I have bred adults from
full-grown larvae taken in nests of the Black-throated Grebe
( Podicipes novae-hollandtae ), which had become stranded upon the
margin of a swamp, and in which the eggs had not yet hatched.
The same nests and, also, small accumulations of drift, i.e ., leaves,
horse- and cow-droppings, etc., blown up upon the margins of
swamp, served as breeding-places for Stomoxys calcitrans and
Sarcophaga sp.
Major E. E. Austen, who kindly examined the specimens of
M. domestica from bush localities in N. Territory and
N. Q., determined the latter as a variety of the typical form.
Evidently the distincion is a very fine one, since recently I have
examined a much longer series than was available to that worker,
and I have compared them and their larvae with typical forms (from
town dwellings) and their progeny, without being able to detect
any variations peculiar to one series.
(0, (4), (6) Identified by Professor M. Bkzzi.
(2) Identified by Major E. E. Austen.
(3) Johnston, T. H. and Bancboft, M. J. Proc. Roy. Soc . Queensland. Vol. XXXI, No. 12.
(5) Johnston, T. H. and Bancroft, M. J. Mem. Queensland Museum. Vol. VII, pt. 1.
1920.
95
NEW TSETSE-FLIES {GLOSSINA) FROM
THE BELGIAN CONGO
BY
Professor R. NEWSTEAD, F.R.S.
AND
Miss ALWEN M. EVANS, M.Sc.
[Received for publication 14 March , 1921)
We have just received from our esteemed colleague, Dr. J. Schwetz,
a small collection of tsetse-flies, captured by him in the Belgian Congo.
Included in this collection are eleven examples (6 <J<£, 5 99) of a new and
hitherto undescribed species, all of them taken in the region of the River
Kwango, on the frontier of the Portuguese territory. In his letter,
dated 20.xii.20, Dr. Schwetz says that he had just visited this region
for the first time, and that in returning down the River Kwango by boat
he came to a region abounding in this species and GL palpalis, R.D.
This new species belongs to the * Fusca Group 11 of tsetse-flies, and is
described below as Glossina schwetzi , sp. n., in honour of its discoverer,
who has devoted long years of research into the bionomics of this important
group of insects, and their relation to human trypanosomiasis.
In a former communication in these Annals 2 attention was called to a
variety or race of Glossina fusca, Walker, from the Belgian Congo, in
which the female armature exhibits a marked deviation from the form
of signum found in typical examples from the Gold Coast. The male
genital armature of the Congo examples also differ in the form of the
harpes from those found elsewhere.
We are convinced, therefore, that we have to deal with a well-marked
race of Glossina fusca : this we have given varietal rank under the name
congolensis , var. n.
GLOSSINA SCHWETZ !, sp. nov.
Hairs of the third antennal segment about one-sixth to one-seventh the
1 width of the segment . Wings of the female with the thickened portion of
the anterior transverse vein darker in colour than the rest . Harpes of the
9 6
male divided into three processes , the proximal process short and spine 4 ike.
Female with signum of the uterus consisting of a single chitinous plate , the
long axis transverse and widest in the distal third .
Male : Length, 10-11*9 mm.; proboscis, 2*3-2*8 mm.; width of head,
3*2-3*3 mm.; front at vertex, 0*6-07 mm. ; wing, 11-12*4 nun.
Fig. 1. Male armature of Glossina schzvetzi , sp. n.
j.r., superior clasper; i.r., inferior clasper ; m.p., median process ; inferior median process ;
t;., vesica ; b harpes.
Female : Length, 12*5 mm. ; proboscis, 3*1 mm.; width of head,
3*5”3'6 mm.; front at vertex, 0*75-0*8 mm.; wing, 13*2-13*6 mm.
Male : Head with the posterior surface ' mouse grey' (Austen) 3 ,
antennal cavity pearl-grey, sometimes with a pale vinous tinge; ocellar
97
spot and frontal stripe unicolorous pale brown ; antennae with the distal
two-thirds of the third segment infuscated, the rest pale buff; outstanding
hairs on third segment, short, about one-sixth to one-seventh the width
of the segment. Proboscis bulb pale translucent buff-yellow, the upper
lateral margins brownish or orange-brown, ventral median suture proxi-
mally, dusky to orange-brown. Thorax with the usual distinctive colour
and markings ; stemo-pleurae more or less infuscated ; scutellar bristles
long. Abdomen: Dorsum of first and second segments light brown;
the remaining segments, together with the lateral margins, dark brown
(‘ mummy-brown,' Austen®), the distal angles either unicolorous with the
rest of the abdomen or slightly paler. Legs : Light or dusky ochraceous-
buff; leg I with the femora infuscated on the upper half of the inner
surface, tips of the last two segments dark brown or black ; leg II with
the tips of the last two segments generally more strongly marked than in I;
leg III with the last two segments entirely dark brown or black, paler
beneath, proximally. Wings with the thickened portion of the anterior
transverse vein scarcely darker in colour than the rest. Genital armature:
(fig i) Harpes (A) with three bi-lateral processes; the proximal process
short and distinctly spine-like ; second and third pairs long, slender, and
only very slightly widened proximally; median process scarcely
projecting beyond the inferior claspers (ic) ; the inferior median
process (mfit) narrowly elongated and projecting backwards considerably
beyond the inferior claspers. Superior claspers (s.c.) bluntly bifid.
Female : Colour and pattern similar to that of the male; but the distal
segment of the abdomen usually paler, and the thickened portion of the
anterior transverse vein of the wing darker in colour than the rest. Genital
armature: (fig. 2) External armature possessing no distinctive characters.
Signum of the uterus consisting of a single symmetrical chitinous
plate of the form shown in the figure, the greatest width being 0*38 mm.
Laterally, it bears a pair of black curved thickenings (a.th), and sometimes
a second pair (p.th) lies behind the anterior pair. That portion of the
plate posterior to the level of the thickenings is much more heavily chiti-
nised than the anterior portion, in which a large transparent space occurs
medianly. The upturned anterior processes (a.p) which are a marked
feature of the signum tend to slight variation, and are much smaller in one
individual than those shown in the illustration.
Belgian Congo: River Kwango, Kasongo Lunda, 24.x. 1920, 2 $9;
River Kwango, near Cuango, 24.x. 1920,4 $$ (' Le soir par terre ') 25.x. 1920
3 99 ; River Kwango, near Kundi, i.xi.1920, 2 <f <J (' Le soir sur chemin ’)
Dr. J. Schwetz.
In its general external facies the male of this species bears a striking
resemblance to that of Gl. tabantformis, West; the female, on the other
Fig. 2. Glossina schwetzi, sp. n. 9 - Signum x c. 155.
a.p.y anterior process; a.tb., anterior thickening; p.tb., posterior thickening.
(Genital fossae not shown.)
hand, owing to the darker colour of the thickened portion of the anterior
transverse vein, might easily pass as a specimen of Gl. brevipalpis, Newst.
Both sexes may, however, be readily distinguished from any other members
of the ' Fusca Group ' by the strikingly characteristic genital armature.
Fig. 3.
Glossina schwetzi, sp. n., A ., antennal fringe x c. 325 ; B third segment of antenna x c. 4c.
Glossina tabanijormis , C., antennal fringe x c. 325 ; A, third segment of antenna X c. 40.
Furthermore, the antenna of Gl. schwetzi can be distinguished from
that of Gl. tabaniformis by the shorter fringe of fine hairs on the
anterior edge of the third segment (fig. 3). The ratio of the length of the
longest fringe-hairs to the greatest width of the segment was determined
99
in four specimens of each of the above-named species. They are tabulated
below :
Gl. sebtvetzi, sp. n. i : 5-9 1 : 6*2 I : 7*1 1:7-5
Gl. tabaniformis . 1 : 37 1 : 4*1 1 : 4-3 1 : 4.3
Comparative Table of the Morphological Characters of the
Male Genital Armature
Gl. schwetzi Gl. tabaniformis
Harpes : Fig. 1 Fig. 1, no. 3
Proximal process ... A stout spine, one-fourth Slender and equal in
length of second. length to the second.
Second process.Slender. The same.
Third process (distal) ... Long, slender and simple. Long £nd strongly bifur¬
cate.
Comparative Table of the Morphological Characters of the
Female Genital Armature
Gl. schwetzi
Gl. tabaniformis
Signum of uterus...
Fig. 2
With a bilateral pair of
Lyriform. No bilateral
black curved thicken¬
ings.
curved thickenings.
Long axis.
... Transverse.
Longitudinal.
QLOSSINA FUSCA var. CONQOLBNSiS, var. nov.
Colour and pattern generally as in typical GL fusca, Walker , but the
wings are usually more heavily infuscated. Harpes of male (fig. 4) with
the proximal process reaching to the tip of the second (serrated) process.
Bilateral portions of the signum of the uterus of the female sub-rotund.
Male : Genital armature (fig. 4). Harpes (A) with the narrow proximal
process reaching to the tip of the second, serrated process; the latter
relatively narrow, and more or less suddenly truncate distally; the third,
a somewhat sickle-shaped appendage, with the distal arm very much
shorter than the proximal one.
Female : Genital armature. In the signum of the uterus (fig. 5) the
width is greater than the length; the bilaterally symmetrical plates forming
the main portion of the signum sub-rotund, and separated medially by
only a very shallow depression.
IOO
The foregoing description, and also the illustrations which accompany
it are based upon fourteen examples, representing both sexes, captured in
i
Fig. 4. (1) and (2) Male armature of Glossina fusca var. congolensis, var. n.
(3) Harpes of Glossina fuse a.
j.r., superior clasper ; i.r., inferior dasper; v. f vesica ; i>., harpes; i.o. t intromittent organ ;
b.i., sickle-shaped process of harpes.
the Katompe, and in the Lomami-Kisengwa districts of the Belgian
Congo by Dr. J. Schwetz.
Fic. 6. Glossina/usca, Walker. ?. Signum x c. 155.
cr., crescentic sclerite ; half of main portion of signum ; g.f., genital fossae ; />., posterior plate,
102
Comparative Table of the Morphological Characters of the
. Male Genital Armature
Harpes :
Proximal process
Serrated process
Distal process ...
G.fusca
Fig. 4, no. 3
... Reaching to the middle
distance of the serrated
process.
... Distally broadly lanceo¬
late ; shaft relatively
broad.
... Distal arm of sickle-
shaped process as long
as the proximal one.
G.fusca vat. congolensis
Fig. 4, no. i, 2
Reaching to the tip of
the serrated process.
Distally suddenly trun¬
cate; shaft relatively
narrow.
Distal arm less than half
the length of the
proximal one.
Comparative Table of the Morphological Characters of the
Female Genital Armature
Gl. fuse a
Fig. 6
Signum
Halves of main portion
Chitinisation
Anterior plate ('*./>.)
Greatest width ...
... Sub-conical. Very mark¬
edly convex, sepa¬
rated anteriorly by
deep V-shaped depres¬
sion extending to nearly
one-third of their
depth.
... Generally complete, the
colour varying from
deep . ochraceous to
black.
Absent.
Average 0*56.
Extremes o*6o, 0*46.
Gl.fusca var. congolensis
Fig. 5
Sub-rotund. Slightly
convex, separated an¬
teriorly by very shal¬
low depression.
Generally incomplete,
ochraceous pctaloid
area (/>.) often sur¬
rounded by consider¬
able almost colourless
area.
Generally present.
Average 0*42.
Extremes 0*44, 0 40
REFERENCES
1. —Newstead, R. (1911). Bull. Ent. Research , Vol. II, pp. 9- 36.
2. —Evans, Alwen M. (1919). Ann. Prop. Med. Parasit ., Vol. XIII, pp. 31-56.
3. — Austen, Ernest Edward (1911). A handbook of the Tsetse-Flics (genus Gbssina).
103
ON A COLLECTION OF PAPPATACI
FLIES ( PHLEBOTOMUS) FROM INDIA
BY
Professor R. NEWSTEAD, F.R.S.,
AND
Major J. A. SINTON, V.C., I.M.S.
{Received for publication 14 March , 1921)
The collection of insects which form the subject of this communication
was made by one of us (J. A. Sinton) in the North-west Province of India,
with the exception of the single capture of a specimen of Phlebotomus
major , Annandale, at Simla. It embraces the records of a very large
number of individuals (664) representing three species and one variety,
and, as the pappataci flies of the region in question have not hitherto
been investigated, it seemed desirable that a complete list of the captures
should be recorded, the subject being a cognate one to that of pappataci
fever.
The occurrence of Phlebotomus sergenti , Parrot, is of much interest,
the species being a new addition to the fauna of India.
In the examination of the long series of examples of Phlebotomus
minutus, Rond., it was found to be impossible to draw any hard and
fast lines for the separation of this species from P. antennatus , Newst., 1
since we found so many intermediate forms ranging between typical
examples of each. It may be recalled that the erection of P. antennatus
was based chiefly upon the remarkably short antennal segments, and the
relatively shorter legs; the male genital armature on the other hand
being absolutely identical with that of typical P. minutus . It seems,
therefore, that the differences in the antennal segments can no longer
be considered as of specific importance ; P. antennatus has therefore been
placed as a variety of P. minutus .
We tender out thanks to Miss Alwen M. Evans, M.Sc., for her assistance
in the preparation and determination of the material.
Phlcbolomus papatasii, (Scop).
In the males twelve examples were found to possess an additional
spine on the inferior clasper of the genetalia; in some instances the
spine was bilateral, in other cases unilateral. One example also possessed
a supernumerary spine on the left superior clasper.
India : North-west Frontier Province. Dera Ismail Khan,
2 33,6 99 , bedroom, bungalow, September and October, 1919; 6 33 ,
7 99 , ditto, 14.iv.1920 to 14.V.1920; 40 33, 52 99 , cowshed, 8 .iii.i 920
to 31.iii.1920 ; 35 33 , 74 9$, ditto, i.iv.1920 to 14.iv.1920 ; 3 33 , 10 99,
ditto, 15.iv.1920 to 15.V.1920 ; 1 3, tents near I.M.S. bungalow, 16.iii.1920 ;
4 33, 11 99 , ditto, 12.iv.1920 to 15.v. 1920. Tank, 1 <J, 5 99, inside tents
with floors sunk 3 feet, 29.viii.1919, i.ix.1919, 16.ix.1919; 2 99, ditto,
6.iv.i920. Bannu, 4 33 , 1 9, bedroom, bungalow, 11.viii.1919 to
12.viii.1919. Idak, Tochi Valley, i 9, mud barracks, 26.viii.1919.
Saggu, near Dera Ismail Khan, 19, stable, 14.iii.1920. Jatta Post,
1 9 , mud barracks, 7.^.1920. Murtaza, i 3, 4 99 , mud barracks,
i.ix.1919 ; 1 (j, 6 99 , ditto, 7.^.1920. Hathala, 1 o, 1 9 , mud barracks,
29.viii.1919 ; 1 9, ditto, 6.iv.ig2o.
Phlebotomus minuius, Rondani.
The specimens showed a marked variation in colour ; every form
intervening between the pale typical examples and distinctly dark or
melanic forms were observed. The left superior clasper in one male
possessed a supernumerary spine. The wings of this species also showed
a marked tendency to variation, both as regards size and contour, and also
in the relative length of the anterior forked vein.
India : North-west Frontier Province. Dera Ismail Khan,
118 <3 <3, 34 99, bedroom, bungalow, September and October, 1919;
6 So, 2 99, ditto, 14.iv.1920 to 14.V.1920 ; 3 S3, 7 9?, cowshed, 8.iii.i920
to 31.iii.1920; 15 S3, 8 99 , ditto, i.iv.1920 to 14.iv.1920 ; 1 3, 3 99 ,
tents near I.M.S. bungalow, 16.iii.1920; 36 33, 13 99, ditto, 12.iv.1920
to 15.V.1920. Tank, 3 33,6 99, tents with floors sunk 3 feet, 29.viii.1919,
i.ix.1919, 16.ix.1919. Bannu, 37 33, 37 99, bedroom, bungalow,
n.viii.1919 to 12.viii.1919. Idak, Tochi Valley, 2 33, mud barracks,
26.viii.1919. Khirgi, 2 99, tents, 30.viii.1919. Hathala, 3 99 , mud
barracks, 29.viii.1919.
i os
Phlebotomus minutus var. antennatus, Newst.
Phlebotomus antennatus , Newst. Bull. Ent. Res., Vol. Ill, p. 365.
The variation in colour from light to dark or melanic forms was similar
to that found in P. minutus, but none possessed the intense lapis-lazuli
colour found in the West African forms. 5
India : North-west Frontier Province. Dera Ismail Khan,
11 < 3 d, 3 99 , bedroom, bungalow, September and October, 1919 ; 2 <$<$,
ditto, 14.iv.1920 to 14.V.1920 ; 2 dd , 1 9 , cowshed, 8.iii.i920 to 31.iii.1920 ;
1 <J, 2 ?$, ditto, i.iv.1920 to 14.iv.1920 ; 1 d , 2 99 , tents near I.M.S.
bungalow, 12.iv.1920 to 15.v. 1920, Tank, 3 99 , tents with floors sunk
3 fept, 29.viii.1919, i.ix.1919, 16.ix.1919. Bannu, 2 dd , 7 99 , bedroom,
bungalow, 11.viii.1919, 12.viii.1919.
Phlebotomus sergenti, Parrot.
Parrot 2 first described this species from Algeria; subsequently it was
found by Marzinovsky 3 at Tiflis, by Sinton in Persia, and by Buxton 4 in
Mesopotamia. Hitherto it does not seem to have been recorded from
India.
India : North-west Frontier Province. Dera Ismail Khan,
1 o, cowshed, 8.iii.i920 to 31.iii.1920 ; 1 d , tents near I.M.S. bungalow,
12.iv.1920 to 15.V.1920 ; 3 dd , 1 9 , stable, Tank Road, 7 miles from
Dera Ismail Khan, 6.iv.i920. Murtaza, i d , mud barracks, i.ix.1919.
Phlebotomus major , Annandale. Records Indian Mus., Vol. V, p. 46
(1910).
India : Simla, i 9, 30.v. 1920.
Colour and general facies resembling those of the female in P. papatasii ,
but the wings are relatively longer and narrower. Position of abdominal
hairs doubtful owing to the rubbed condition of the dorsal surface,
but the hairs on the venter more or less erect. Palpi with the 2nd, 3rd,
and 4th sqjments sub-equal in length ; the 5th twice the length of the 3rd.
Total length of head and body, 3*5 mm. ; length of wing, 3*3 ; greatest
width, 1-05 mm. ; length of leg ii, 4-8 mm. ; femur more than half the
length of the tibia, the latter equal in length to the first three proximal
segments of the tarsus.
This, as may be gathered from the above details, is an exceptionally
large species, and in its general facies and certain structural characters
agrees with Annandale's description of his P. major {lx.). There are some
io6
important discrepancies however : i.e., in the arrangement of the abdomi¬
nal hairs, and the length of the body as compared with that of the legs.
We feel, however, that these may be considered as possible mutations
within the range of variation of the species, and have therefore placed
it here.
REFERENCES
1. —Nkwstead (1912). Bull, Ent. Res., Vol. Ill, p. 365.
2. —Parrot (1917). Bull. Soc. Path. Exot., X, p. 564.
3. —Marzinoysky (1917). Medical Review , Moscow, LXXXVII, p. 612.
4. — Niwstead (1920). Bull . Ent. Res., Vol. XT, p. 307.
5. - (1920). Bull. Ent . Res ., Vol. XI, p. 305.
107
NOTES ON ORIENTAL SORE IN RUSSIAN
TURKESTAN AND THE RESULTS OF
TREATMENT WITH INJECTIONS OF
TARTAR EMETIC SOLUTION
BV
J. A. SINTON, M.D., Major, I.M.S.
(i Received for publication 27 April , 1921)
The cases reported below seemed worthy of record on account
of the rapidity with which they reacted to treatment with intravenous
injections of solutions of tartar emetic, a treatment which does not
seem tp have been tried previously in Turkestan.
This disease is very widely spread in Russian Turkestan, and
is said to be found in all the towns on* the Trans-Caspian Railway
from Askhabad to Tashkent. Yakimoff and Schockov (1915)
described it at Askhabad, Boukhara, Samarkand and Termfcze, and
in the latter place it seemed especially prevalent.
Locally it is said that practically every person residing in
Turkestan for five years has the marks of at least one of these sores.
At 'Tedjen station, on casual inspection, about 50 per cent, of the
people seemed to have marks of old sores on their faces, but the
authors quoted above state that only 58 2 per cent, of the sores
examined by them were leishmaniasis. The same authors report
two cases of cutaneous leishmaniasis in dogs in Turkestan.
In all the six cases noted below LeisJimania tropica were found.
The treatment used was intravenous injections of 2 per cent,
solution of antimonium tartaratum (tartar emetic) in normal saline
solution; the sores received no special local treatment. Except for
a little vomiting immediately after the injection, in two cases, no
constitutional effects were caused by the injections.
io8
NOTES ON THE CASES
Case i. Russian Nursing Sister at Kaakha, Turkestan.
History . Resident in Turkestan for some years. Duration of sores about
three months.
Condition, (a) A small raised nodular sore J-inch in diameter at the angle
of the left jaw, with slight ulceration in the centre, (b) Three similar sores on
the left forearm.
Treatment. 5.10.18. Intravenous injection of 3 c.c. T.E. solution.
7.10.18. Injection of 4 c.c. T.E. solution.
Result . On account of military operations the case was not seen again until
14.11.18, at which time all the sores were healed and the patient reported that
they had all healed within three weeks after the injections.
Case 2. Sepoy A.S.
History . Patient has been in Turkestan about three months. Duration of
lesion uncertain.
Condition. Sore about £-inch diameter on right hand. Warty and ulcerated.
Treatment. 19.1.19. Intravenous injection of 2 c.c. of T.E. solution.
22.1.19. Injection of 3 c.c. of T.E. solution.
Result. Completely healed by 27.1.19.
Case 3. Sepoy M.K.
History. Had a small abrasion on the back of the left hand about 1.10.18,
at Meshed, N. Persia, which gradually got bigger. He arrived in Turkestan
on 10.10.18.
Condition. On 7.1.19, at Bairam Ali, Turkestan. A large sore about i£ inches
in diameter on back of left hand with inflamed margins and a warty base. No
enlarged glands or thickening of the lymphatic vessels.
Treatment. 14.1.19. Injection of 2 c.c. of T.E. solution.
16.1.19. » » 3 c.c. „ „
I 9- I - I 9* » » 4 c.c. „ „
22.1.19. „ „ 5 c.c. „ „
Result . The warty growth gradually shrivelled and the ulceration
disappeared. Completely healed on 28.1.19.
Case 4. Sepoy S.S.
History. Got an abrasion of the left hand at Dushak, Turkestan, on 14.10.18,
which gradually increased in size. Duration three months.
Condition, (a) A large circular lesion about inches in diameter on back
of left hand with infiltrated edges and a warty base with slight ulceration in the
centre. (< b ) A few small non-ulcerated nodules in the adjacent skin.
Treatment. 14.1.19. Injection of 2 c.c. of T.E. solution.
16.1.19* » 99 3 C, ^ # 99
19- 1 l 9 - 99 »4 cc - » „
21.1.19* » » 5 cc - » »
Result. After the second injection, a firm nodule the size of a pea was
discovered in a lymphatic vessel about zl inches below the elbow at the back of
109
the left forearm, and a soft swelling about 2 inches by I inch on the inner side of
the same forearm. This swelling was on the opposite side to that on which the
injections had been given. Marked improvement was noted at the time of the
third injection, the sore being dry in the centre and the warty growth shrivelling.
The sore was completely healed on 29.1.19.
Case 5. Sepoy C.D.
History . Said to be of two weeks’ duration. No history of wound or abrasion.
Patient has been several months in Turkestan.
Condition. At the beginning of January, 1919.
( a ) A circular lesion with thickened edges and a warty base about 1 inch in
diameter on front of left wrist.
( b ) Two small nodules in the lymphatic vessels about two inches above
the sore.
Treatment. 14.1.19. Injection of 2 c.c. of T.E. solution.
16.1.19. » » 3 c c - >> »
19.1.19. „ „ 4 c.c. „ „
22.1.19* » » 5 c,c * » »
Result. The warty growth rapidly shrivelled and the nodules disappeared
from the lymphatics. Completely healed on 27.1.19.
Case 6. Sepoy N.S.
History. He has been in Turkestan four months. He states that the lesions
began with great itching and appeared more or less simultaneously. Duration
about one month.
Condition. On 1.1.19.
(a) On the front of the left wrist was an oval sore about 1 inch by $ inch,
with thickened edges and a warty centre, but no ulceration.
(b) On the front of the right forearm :
(1) A circular lesion, 1 inch in diameter, with heaped-up and
thickened edges, having an ulcer J~inch in diameter in the
middle with a granulating base.
(2) A small sore J-inch in diameter, with a thickened raised edge
and a minute ulcer in the centre like a boil.
(3) There were two small nodules in the lymphatic vessels about
two inches below these sores, and there were no enlarged glands.
Treatment. 16.1.19. Injection of I c.c. of T.E. solution.
i* 1 - 1 * » » 3 c.c. „ „
22.1.19. „ „ 3 c.c. „ „
Result. At the time of the first injection some of the solution passed under
the skin because the patient suddenly moved his arm. At the time of the second
injection there was marked thickening and some inflammation at the site of the
previous injection. Improvement was noticed in the sores on 22.1.19, and this
was marked on 25.1.19. There was still some infiltration of the area around the
point of first injection on 25.1.19, but there was no suppuration. Completely
healed on 30.1.19.
no
These cases may be summarised as follows: —
Table I.
Case
No.
Duration
before
treatment
Amount
of T.E.
injected
Number
of
injections
Duration of
treatment
in days
Number of
days till
cured.
X
3 months
cgms.
■ 4
2
3
? 21
2
?
1
IO
2
4
8
3
34 months
1 28
4
9
14
4
3 months
28
4
9
*5
5
4 month
28
4
9
>3
6
1 month
14
1
3
7
>4
Average
1
2-2 months
zo -33
3 -i
6-8
14* 1
An analysis of the clinical signs in these six cases shows that
Cases 2, 3, 4, 5 and 6 correspond to the ‘ cutaneous hypertrophic
non-ulcerating papillomatous 9 type of da Matta’s classification
(1916) of the leishmaniases, but in this type in these Turkestan cases
it was found that after a certain time these warty growths tend to
ulcerate in the centre. As they bleed very easily, this ulceration
seems most probably due to injury followed by a secondary infection.
In Cases 1 and 6 the typical ‘ Oriental Sore * of the text-book
was found.
In Cases 1, 4 and 6 the lesions were multiple, and in Case 6
both the papillomatous type and the typical oriental sore were found.
Yakimoff and Schockov (1915) state that a single sore is commonest,
but as many as seventeen may occur.
In Cases 4, 5 and 6 nodules were present in the lymphatic
vessels within a few inches of the sores, those in Case 6 being
remarkable in that they were distal to the lesions.
The attached table gives a comparison between the results of
treatment with intravenous injections of tartar emetic solution only,
in the cases reported by myself (1917), in those reported by Greig
(1917) in the same year, and in the present cases.
Ill
Tabli II.
•
, Sinton, 1917
Greig, 1917
Sinton, 1919
Where contracted.
N.W. Frontier,
India and
Mesopotamia
Mesopotamia
Russian Turkestan
Number of cases treated.
6
iS
6
Average duration of the disease
before treatment .
2 months
2*6 months
2*2 months
Duration of the treatment in
days.:—
-
Average .
20
6-8
Maximum.
37
...
9
Minimum.
8
...
3
Number of days from commence¬
ment of treatment till cured :—
Average .
28-3
28*2*
! 4 ‘i
Maximum.
5 1
52 #
21 (?)
Minimum.
12
i6 #
8
Amount of T.E. given
intravenously!—
Average .
38 2 cgms.
71*5 cgms.
20-33 cgms.
Maximum.
89 cgms.
150 cgms.
28 cgms.
Minimum.
12 cgms.
20 cgms.
10 cgms.
Number of injections given :—
Average .
5'3
?
3
Maximum.
9
?
4
Minimum.
3
?
2
Result
Cured .
6
*7
6
Not cured .
0
1
0
•Note.—T hese figures are probably excessive, as they represent the ‘ number of days
under observation in hospital/
112
From this table it will be seen that the amount of tartar emetic
needed to produce a cure in the Turkestan cases was only half to
one-third of that needed in the other cases, and the rate of cure
was almost twice as rkpid.
Although it is not possible to draw any definite conclusions from
so few cases, yet it would seem probable that the variety of sore
found in Turkestan is more amenable to treatment with antimony
than the Mesopotamian type.
I am indebted to Dr. Minkavitch, of the Russian Medical
Service, for the following details of the treatment adopted for the
cure of this disease by some medical practitioners in Turkestan.
This treatment is a modification of the local native treatment.
A piece of Emplastrum Cantharidis is cut slightly larger than
the sore; this is placed over the sore and completely covered by a
larger piece of adhesive plaster to hold it in position. This dressing
is renewed daily, and the blisters which have formed are opened.
This treatment is continued for four days, after which the sore is
treated with some simple ointment. Some very good results are said
to be obtained by this treatment, but the scar left is large.
REFERENCES
Da Matta, A. (1916). Tableau synoptique de la classification des Leishmaniotes. Bull. Soe.
Path. Exot., Vol. IX, pp. 101-102, quoted in Trop. Dis. Bull., Vol. IX, p. 231.
Greig, E. D. W. ( 1917 )- Summary of the results of the observations on the treatment of
Oriental Sore by Antimonium Tartars turn. Ini. Journ. Med . Res., Vol. V, pp. 394-400.
Sinton. J. A. (1917). The treatment of cutaneous Leishmaniasis with intravenous injections of
tartar emetic. Ind. Med. Gaz ., Vol. LIT, pp. 239-241.
Yakimoff, W. L., and Schockov, N. F. (1915). Leishmaniose cutanee (bouton d'orient)
au Turkestan Russe. C. R. Soc. Biol., Vol. LXXVIII, pp. 107-109, quoted in Trop.
Dis. Bull., Vol. VI, p. 225.
A METHOD FOR THE CULTIVATION
OF BLASTOCYSTIS
BY
HARVEY P. BARRET
From the Laboratory of The Charlotte Sanatorium ,
Charlotte , N.C.
(Received for publication 13 April , 1921)
Quite a good deal of uncertainty has existed as to the true nature
of blastocystis. Prowazek (1904) and Ucke (1907) first described
this organism as a cyst of trichomonas. Alexeieff (1911) was the
first to describe blastocystis as a distinct organism of a vegetable
nature. Others have described this organism and discussed the
possibility of its being a cyst of trichomonas or other intestinal
parasite, or some degeneration form of these parasites.
Bohne and Prowazek (1908), Benson (1909), Wenyon (1910),
Prowazek (1911), Brumpt (1912), Chatton and Lalung Bonaire
(1912), Macfie (1915), Swellengrebel (1917), Chatton (1917), have all
given various descriptions of this organism. Wenyon (1920) gives a
very comprehensive review and digest of the subject. Parts of his
article are quoted below : —
‘ What then is the blastocystis which occurs so commonly in the
human intestine and that of other animals? Prowazek (1911)
maintained that they were cysts of trichomonas, but there is no
evidence to support this view. Swellengrebel (1917) has suggested
that they are degenerate forms of various intestinal protozoa, while
Jepps and Dobell (1918) have noted that certain degenerate forms
of Dientamoeba fragilis resemble dead blastocystis. I myself have,
for want of evidence to the contrary, always regarded them as of a
vegetable nature, and this may be the case in spite of the resemblance
to the cysts of Prowaeekella lacertae. Swellengrebers conclusion is
that blastocystis " is not the name of a zoological genus but of a
peculiar form of degeneration to which representatives of different
genera of intestinal protozoa may be liable.” 9 Wenyon concludes,
‘ It is evident, therefore, that there is a difference of opinion
as to the true nature of blastocystis, and we must await further
information.*
The writer has had under observation for the past few years a
case of balantidial infection, whose stools are loaded with blasto-
cystis as well as balantidia. While attempting to cultivate the
balantidia, the blastocystis was found to multiply readily on
the medium used. Since then numerous cultures from different
individuals have been made. This work was begun in 1919, and
the first successful culture made in September of that year. The
universal occurrence of this organism may be noted from the articles
by Lynch.(i9i7) in the United States, Wenyon and O’Connor (1917)
in Egypt, Haughwout (1918) in the Philippines, and Maplestone
(1921) in Australia.
METHOD
The culture medium used in this work is very simple, being
made up of human blood serum and 0*5 per cent, of sodium chloride
solution. Various concentrations of the serum in salt solution have
been tried. The most favourable strength has been 10 per cent.
Walker (1913) found 0 5 per cent, salt solution to be the propef
tonicity for Balantidium colt , and this concentration has been used
in the work with blastocystis. The salt solution is sterilized in the
autoclav and the serum added after inactivation at 55 0 C. for one
half-hour. The pooled serum of several individuals has been used
instead of that from a single individual, although no work has been
done to show whether or not one serum may be inhibitory while
another is favourable to the growth of blastocystis. The sera of
animals other than man have not been tried. TTiis medium is
faintly alkaline to litmus. The medium is distributed in narrow
test tubes in quantity sufficient to give a column of fluid at least
100 mm. high.
No growth takes place at the surface of the tube, and the parasites
multiply best at the lower portion of the tube, evidently needing
little free oxygen for their growth.
In making the initial inoculation, a small portion of stool or an
emulsion of faeces in salt solution is placed at the bottom of the
tube containing the culture medium. The culture is incubated at
37 0 C. for twenty-four to forty-eight hours and then examined for
blastocystis. It is best to examine early cultures every twenty-four
hours in order to note the degree Of growth of blastocystis and to
make transfers before bacterial contamination is too heavy. If good
growth is obtained after twenty-four hours, a transplant is then
made into fresh culture medium, using the sediment in the original
tube. When the organisms have established themselves in the new
medium, it is best to make transfers every forty-eight hours.
Cultures seventy-two hours old, or older, will hardly ever give good
growth in succeeding transfers. As stated above, the optimum
growth takes place in the lower portion of the tube. In making all
subsequent transfers the sediment from the cultures is used, and is
introduced into the bottom of the tubes of the fresh medium. Using
this method, the writer has carried blastocystis through more than
twenty-five sub-cultures, covering a period of about fifty days.
Cultures have been abandoned, usually because of pressure of
other work or because of accident or Carelessness in making
transfers, although they often die from bacterial overgrowth. With
proper care a culture can probably be carried on indefinitely.
The writer has made no attempt to classify different strains of
blastocystis though, the gross appearance of organisms from
different individuals would suggest strongly that several strains of
blastocystis have been encountered. Nor has any attempt been
made to trace out a developmental cycle for blastocystis. This task
is properly left to one proficient in systematic mycology.
In cultures, budding, as described by Alexeieff, and binary
division are the methods of multiplication exhibited by this
organism; both of these processes have been observed in all cultures.
In the examination of hundreds of preparations the so-called multiple
division form of Alexeieff, pictured by Wenyon and O’Connor (1917),
has been encountered on only two occasions. This form would seem to
be some freak in the development of the organism rather than a true
developmental phase. In cultures, as in the intestinal contents, a
great variation in the size of the organisms is observed. It may be
said with a reasonable degree of certainty that the large vacuolated
forms are the result of degeneration, as these forms are commonly
found in old cultures, and successful transfers cannot be made from
cultures made up of these forms. No flagellate forms, 'and no forms
showing amoeboid or other motion have been encountered. Blasto¬
cystis retains its original form after twenty-five successive transfers
ii 6
in artificial culture medium, and no forms were seen in the twenty-
fifth transfer that were not seen in the first.
This work is submitted as proof that blastocystis is a distinct
zoological genus, and not a cyst of protozoa nor a form assumed by
protozoa undergoing degeneration.
REFERENCES
Alexeietf, A. (191 r). Sur let 1 Kystes de Trichomonasintestinalis * dans l'intestin dec Batradens.
Bull . Sci. France et Belgique , Vol. XLIV, p. 333.
- (1911). Sur la nature det formations dites 4 Kystes de Trichomonas intestinalis.*
C. R. Soc. Biol ., Vol LXXI, p. 296.
Benson (1910). Untersuchungen fiber Trichomonas intestinalis und vaginnlis des Menschen.
Arch. /. Protutenk. y Vol. XVIII, p. 116.
Bobne und Prowazek, S. (1908). Zur Frage der Flagellatendysenterie. Arcb.f. Protistenk^
Vol. XII, p. 1.
Brumft, E. (1912). Colite i Tetramitus mesnili (Wenyon 1910) et colite I Trichomonas intestinalis
Leuckart 1879. Blastocystis bominis , n. sp. et formes voasines. Bull. Soc. Path. Exot n
VoL V, p. 725.
Chatton, E. (1917)* Let 4 Blastocystis' stades du cycle 6volutif de flagelles intestinaux.
C. R. Soc. Biol., VoL LXXX, p. 555.
Chatton, E. it Lalung-Bonnaire (1912). Amibe Umax {Vahlkamfia n. gen.) dans l'intestin
humain. Son importance pour I'interprttation des amibes de culture. Bull. Soc.
Path. Exot.y Vol V, p. 135.
Haughwout, F. G. (1918). The tissue invasive powers of the flagellated and dUated protozoa,
with etpedal reference to Trichomonas intestinalis. A critical review. Philippine
Joum. of Science , Sec. B, VoL XIII, p. 217.
Jeffs, M. W., and Dobell, C. (1918). Dientamoeba fragilis n.g., n. sp., a new intestinal amoeba
from man. Parasitology , Vol. X, p. 352.
Lynch, K. M. (1917)* Blastocystis bominis ; its characteristics and its prevalence in intestinal
content and faeces in South Carolina. Journal of Bacteriology , Vol. II, p. 369.
Machs, J. W. S. (1915)* A case of dysentery in a monkey in which amoebae and spirochaetes
were found. Ann. Trop. Med. and Parasit ., Vol. IX, p. 507.
Maplestone, P. A. (1921)* Human Intestinal Protozoa in North Queensland. Ann. Trop.
Med. and Parasit ., Vol. XIV, p. 283.
Prowazek, S. (1904). Untersuchungen fiber einige parasitische Flagella ten. Art. a. d. Kais.
Gesundbeitsamte , Vol. XXI, p. 1.
-(1911). Zur Kenntnis der Flagellaten des Darmtraktus. Arch, fur Protutenk.,
VoL XXIII, p. 96.
Swellengrebel, N. H. (1917)- Observations on Blastocystis bominis. Parasitology , Vol. IX,
p. 451.
Walker, E. L. (1913). Quantitative determination of the balantididdal activity of certain
drugs and chemicals as a basis for treatment of infections with Balantidium coli.
Philippine Journ. of Science , Sec. B, VoL VIII, p. 1.
Wenyon, C. M. (I910). A new flagellate, 1 Macrostoma mesnili ' n. sp., from the human
intestine, with some remains on the supposed cysts of 4 Trichomonas' Parasitology,
Vol. Ill, p. 210.
-- (1920). Observations on the intestinal protozoa of three Egyptian lizards, with a
note on a cell-invading fungus. Parasitology , Vol. XII, p. 350.
Wenyon, C. M., and O'Connor, F. W. (1917)* Human intestinal protozoa in the Near East.
Wellcome Bureau of Scientific Research , London.
"7
THE INCIDENCE OF INTESTINAL PARA¬
SITES, ESPECIALLY WITH REGARD TO
THE PROTOZOA, AMONGST SYMPTOM¬
LESS CARRIERS IN JAMAICA.
BY
HENRY HAROLD SCOTT,
M.D., M.R.C.P. (Lond.), F.R.S.E., D.P.H.
GOVERNMENT BACTERIOLOGIST, HONG KONG; LATE GOVERNMENT BACTERIOLOGIST,
JAMAICA, B.W4.
(Received for publication 13 April , 1921)
The objects of this investigation were, firstly, to find out the
incidence of infection with various species of intestinal parasites in
patients admitted to hospital for conditions totally unconnected with
dysentery or other intestinal complaints, i.e ., symptomless carriers,
and, secondly, to see what strains of amoebae were common.
That three successive examinations should indicate freedom from
infection was a purely arbitrary standard, and is shown later to be
erroneous in some at least of the cases in this series.
We must bear in mind that a ‘ negative examination * does not
mean that no cysts are present; it, of course, merely implies that
none have been found, and, when we consider how small a proportion
of the whole stool is submitted to examination, we can understand
that if the cysts are few in number they may easily be missed in
such a limited examination. Again, if only one or two are found
after thorough examination of several specimens, though the results
will be erroneous from the statistical point of view if put down as
negative, owing to none being found in the first three specimens,
nevertheless the infection is probably so slight that for all practical
purposes the stool is negative.
In this connection, however, it is also important to remember that
the value of a negative report on examination will differ somewhat
according to the species of parasite. Thus, a negative examination
in respect of Giardia intestinalis is more likely to be correct, as
Ii8
regards inference, than one in respect of Entamoeba coli t owing to
the more general and even distribution of the former in a stool;
while the latter are often so irregularly distributed through the faeces
that one sample taken for examination is by no means representative
of the whole stool. Under conditions such as the latter, two
observers examining different portions of the same stool might give
diametrically opposite reports: the one that he could not find
any, the other that coli cysts were abundant. In cases of Giardia
infection this is less likely to occur.
In order to minimise this difficulty as far as was practicable,
instead of taking these specimens from a stool and examining direct,
1 have adopted the concentration method of Cropper and Row
(1917). By this means, not only was the examination facilitated,
but one was able to employ larger quantities for examination, and,
by the prolonged shaking and consequently more thorough
emulsification, to obtain specimens more representative of the general
state of the stool.
A second branch of investigation was also undertaken, as the
opportunity to add something to the knowledge of the size of cysts
in different persons infected was too good to be lost. All who have
undertaken the measurement of a large number of cysts of the various
parasites will appreciate the tediousness and labour involved in
doing this day after day, but it appeared to me that it would be a
matter of interest and possible utility to see whether the researches
of Matthews and Malins Smith (1917) in Liverpool cases were
corroborated by the findings in patients in Jamaica.
With a pressure of routine work, and having no trained
assistants, I have been able to deal with a limited number of cases
only, very limited indeed as compared with the large numbers
examined and reported upon by the two authors mentioned, and
in putting forward my results, I do so with a feeling of diffidence,
since, owing to the limited time and resources at my disposal, I do
not intend them as a comparison with their brilliant work.
The remarks which follow, however, will serve as a good basis
for further investigation which can be continued at any time, and
also as a basis for comparison with similar cases in other tropical
countries.
The method which has been employed is as follows :—Cases for
examination were selected from among patients in the hospital to
which the laboratory is attached. Those patients who had been
admitted for any intestinal condition were excluded. It was
desired to find out whether any of those admitted to hospital for
complaints other than intestinal were harbouring parasites. Since
these patients would leave hospital as soon as they recovered, they
would then become healthy carriers, as regards the intestinal
parasites present.
A certain number of male and female patients were selected, and
fresh stools from them were sent to the laboratory on alternate days
for three weeks; on the intermediate days a similar series of an equal
number of patients was sent. Two series of patients could thus be
dealt with at the same time, or rather during the same period, the
stools from one series being brought to the laboratory on Mondays,
Wednesdays and Fridays, and those from the other on Tuesdays,
Thursdays and Saturdays. Since no specimens were sent up on
Sundays, each case was examined on the first, third, fifth, eighth,
tenth, twelfth, fifteenth, seventeenth, and nineteenth days.
As none of the patients showed any signs of intestinal infection
they were not under any treatment for the condition even when
protozoal infection was detected, so that the natural course and
variations could be followed during the period of examination.
As Dobell (1917) has stated: ‘A minimum of six examinations
per case should be adopted (it is an arbitrary number), no case being
regarded as free from infection [he is speaking of Entamoeba
histolytica ] unless it has been examined six times with negative
results. Untreated cases may be examined on any six days
convenient to the examiner, since there is no evidence of periodicity
in the occurrence of positive or negative examinations. Since the
distribution appears to be at random, the chances of finding the
infection, if present, are as great for any one day as any other.’
The case J. T. affords an example in which infection with
E. histolytica was not detected until the last examination of all; six
examinations would have failed to find this one.
The period of three weeks being decided upon, each patient’s
stools, if the actions were regular, were examined on nine occasions.
Cropper and Row’s concentration method was adopted, and was
found to facilitate matters greatly when later we came on to the
120
mensuration of the cysts. There is no need to detail the method;
suffice it to say that after concentration three separate specimens
were examined from each before the stool was pronounced negative
for the day.
Examination for helminth ova was also made on the first day
that the stools were sent; if these were found, no further search was
made for them on subsequent occasions, as the percentage of cases
infected with these ova has been given in detail in my reports from
Jamaica for several years, totalling over 40,000 specimens. If,
however, none were found on the first day, their presence was noted
when one came across them later in searching for protozoal cysts.
The tables appended show the distribution of the various
parasites found, the cases being arranged according to age. It will
be seen that the stools from one hundred and two patients have been
submitted to examination, of which fifty-three were males and
forty-nine females.
In twenty-one of the forty-nine female cases the Entamoeba colt
was found during these examinations, and in six the E . histolytica.
It will be noticed that in only one of these was the latter found
without the former. Cysts of Giardia intestindlis were found alone
in four instances, and in combination with other forms in eleven.
Chilomastix mesnili was met with seven times and Balantidium colt
once only.
Comparing these figures with the numbers found among the
fifty-three male cases: E. histolytica was found in nine, E . coli in
twenty-seven, G. intesiinalis in twenty-one (in four of these they
were present alone), C. mesnili was found in six, and B. coli in one.
Thus, out of the total of one hundred and two patients whose
stools were examined, E . coli was found in forty-eight, or just over
47 per cent.; E. histolytica in fifteen, or 14*7 per cent.; G. intesiinalis
in thirty-six, or 35*29 per cent.; C. mesnili in thirteen, or 1274 per
cent.; and B . coli twice.
Cercomonas sp. occurred fairly frequently, but no record was kept
of this. It is seen in a considerable number of the ordinary routine
specimens sent to the laboratory for examination for ankylostome
ova.
These proportions, when we consider that none of the patients
gave any history, at all events recent, of intestinal troubles, are very
121
high, and it was my intention to continue the investigation to see
whether subsequent series maintained this high percentage, but my
transfer to Hong Kong has prevented this.
Before considering each of these protozoa in greater detail, it
will be well to note the criteria made use of in distinguishing the
coli and the histolytica cysts; those of Giardia and Chilontastix need
no description. As regards the points relied upon in differentiating
coli from histolytica cysts, as is well known, it is the general
summation of various characters which enables a definite diagnosis
to be made in almost every case; no single character considered alone
will suffice for a decision. The points are: —
1. Size.
2. Nuclei: ( a) number, (b) character and arrangement of the
chromatin.
3. Cytoplasm.
4. Inclusions: ( a ) chromatoid bodies, (b) vacuoles.
5. Cyst wall.
6. Shape.
1. * Sise. The vast majority of histolytica cysts have a diameter
between 5/1 and 15 p, and of these there are said to be two chief
strains met with, namely, ‘small,’ between 6/1 and gp, with an
average diameter of 7*7/*, and the ‘ ordinary,’ between lOfi and 14/1,
with an average of 12*6/1.
• Coli cysts vary between wider limits, namely, ii/u to 35/1, the
commonest being between 14/u and 22ft, so that difficulties under
this head are only likely to arise in differentiating between a large
histolytica and a small coli cyst.
2. Nuclei : (a) Number. In histolytica they may vary from one
to four,. Malins Smith states that more than half are quadrinucleate,
while one-third are uninucleate. In contradistinction to this, more
than four-fifths of coli cysts contain eight nuclei, and only about
10 per cent, are quadrinucleate, or less; occasionally a cyst with
sixteen nuclei may be met with. It would appear that the
consistence of the stool has some influence; thus, it is stated that in
loose stools about 60 per cent, of the cysts contain eight nuclei, in
semi-formed between 80 and 90 per cent., and in formed stools
between 90 and 95 per cent. Binucleate cysts are found in about
7 per cent., and quadrinucleate in 3 per cent.
122
(,b ) Character and arrangement of the nuclear chromatin. The
situation of this is, in either case, peripheral, but in histolytica it
consists of smaller granules and is distributed fairly evenly, whereas
in colt it is in larger masses or small blocks less evenly distributed,
and therefore outlines the nucleus more definitely than is usually the
case in histolytica.
3. Cytoplasm. This is of a greenish hue in a fresh specimen,
of histolytica and is not uniform in appearance, whereas in coli the
uniformity is greater, the colour is pale and more of a greyish tint,
4. Inclusions : ( a ) Chromatoid bodies. These are. better seen
in fresh saline specimens than in preparations put up with WeigerVs
iodine. In histolytica they are found in about 30 per cent, of the
cysts, are rod-shaped, with blunt, square or rounded ends; whereas
they are less commonly seen in coli , only about one in twenty
containing them, they are more irregular, are often pointed at the
end, or ‘splintered.’ The value of chromatoid bodies in diagnosis
is variable, for they may be absent from two-thirds of the cysts.
( b ) Vacuoles. These are fairly often seen in histolytica , may be
multiple, and usually have an ill-defined contour and stain faintly
with iodine; in contradistinction the vacuoles of coli are rarely
multiple, have a sharply defined edge, and stain more deeply with
iodine. —r
5. Cyst wall. This is usually thinner in histolytica than in
coli , and the latter sometimes appears to have a thin second contour;
these points, however, are far from distinct, and are of minor
importance as aiding diagnosis. ... ; .
6. Shape. Coli is stated to be less frequently asymmetrical than
is histolytica , but one comes across cases in which quite a
considerable proportion of the coli cysts are asymmetrical; this was
particularly noticed in one of the patients in my series.
When the majority of these points are in evidence there is no
difficulty in diagnosis, but one may meet with cysts which have two
or four nuclei, are ill-defined as to colour and uniformity of the
cytoplasm (as stated by Matthews and Malins Smith), have no
chromatoid bodies, no vacuoles, and not a sufficiently distinctive
arrangement of the peripheral chromatin; in such cases certain
diagnosis is not possible, but fortunately these are rare.
123
ENTAMOEBA COL I CYSTS
These were present, as already stated, in 47 per cent, of the
one hundred and two cases examined; there was a slightly larger
proportion found among males, viz., twenty-seven out of fifty-three
(in the females twenty-one out of forty-nine). Since at least five
hundred cases should be examined before any reliable inferences can
be drawn as to incidence, it is very probable that further examina¬
tions would alter this preponderance. As shown in the table, it was
the only one found in five cases of the males and in eight of the
females; in conjunction with others it was more often met with. In
some instances the infection was multiple; thus, in two of the males
and one of the females there were Ankylostotna , Ascaris, Trichiuris %
coli , histolytica , Giardia and Chilomastix , without causing any
apparent abdominal or intestinal disturbance.
No age seems to be exempt; thus, the cysts were found in a boy
of 9 years of age and in a man of 68 years, in a girl of 13 and a
woman of 56. None were found in the five women whose ages
exceeded this, probably a mere coincidence.
It is interesting to remark that the nearest approach to the
proportion of coli infection found in this series of natives examined
(namely, 47 per cent.) was noted by Matthews and Malins Smith in
England among asylum patients (45 9 per cent.). For those who
are acquainted with the West Indian native, the obvious inference
would form a subject of acrimonious if not fruitful debate.
Mention has already been made of the irregularity of distribution
of the cysts of coli in the faeces, resulting in the finding of several
in one specimen and none in others in the same stool; but, by
examining larger amounts and concentrating, as I tried to do, the
effects of this irregularity have been reduced. The fact must also
be borne in mind that the cysts may appear intermittently and be
found in nearly all the preparations made one day, but absent on
another in spite of the careful examination of many preparations.
Several of the cases given in the table bear out this point. Thus, in
No. 52 of the males no cysts were found until the middle of the
third week of examination, and in No. 16 they were seen on the first
day in conjunction with histolytica , but not again, although the
latter was found once subsequently. Other instances of intermittency
124
among the males are Nos. 4, 6, 15, 18, 49. More cases of a single
finding of this variety of cyst occurred among the women, namely,
Nos. 18, 25, 9, 41, 27, 3, when they were seen at the first, third,
fifth and sixth examinations only. In Nos. 42 and 24 they were not
seen until the seventh and eighth examinations respectively, and,
although there was infection in No. 22 with coli, histolytica , Giardia
and Chilomastix , the first-named was found on one occasion only.
It has been stated by Dobell that ‘ in a series of cases in which
the mean number of examinations per case is three, not more than
two-thirds, and possibly not more than half, will be detected/
Taking the first three examinations in this series of cases, there were
detected fourteen among the females and eighteen among the males,
a total of thirty-two out of the forty-eight in whom these cysts were
present—exactly two-thirds.
Matthews and Mai ins Smith found that six examinations give for
histolytica about three times the number of positive results obtained
by one examination (23 per cent, in place of 7*8 per cent.), and for
coli 20*6 per cent, from one examination became 59*8 per cent, by
subsequent ones.
In my series, five only of the twenty-two positive cases among
the females were detected at the first examination (less than one in
four), and ten of the men, giving a total of fifteen out of forty-eight
in spite of the concentration methods.
BNTAMOBBA HISTOLYTICA CYSTS
The cysts of histolytica are usually more regularly distributed
than those of coli, though less so than those of Giardia . A negative
record may, as Dobell states, arise either from their having a localised
distribution and the part containing them not being submitted to
examination, or to their being few in number and so overlooked. He
offers an explanation as to the uniformity or localisation of distribu¬
tion in the site of infection; if the upper part of the large intestine
is affected, the faeces here being more or less fluid the cysts are fairly
evenly distributed, and when solidification of the faeces occurs lower
down the cysts remain scattered through the mass, but, if the lower
part is affected, the faeces are already practically solidified and the
cysts from the ulcerated surface are thus more superficially distributed
125
and also localised to those parts of the contents which have been
in contact with the infected, ulcerated surface.
There would seem to be no periodicity in the appearance of cysts
of histolytica in the faeces. Owing to the scarcity or to the localised
distribution a negative record must not be taken to mean non¬
infection, unless a large number of examinations are made. Three
negative examinations certainly do not warrant a negative report.
Thus, only three of the cases among the females were discovered in
my series in the first three examinations and not a single one at the
first, in spite of the concentration method, showing most emphatically
the uselessness of a negative single finding. Among the males only
two were found at the first examination, and only six in the first
three. Of the men and women together, only nine out of fifteen were
discovered in the first three examinations. At the fourth one more
was found, another at the fifth, three more at the sixth, and one not
until the last time of examination.
The tables show also the irregularity in the days of appearance
of the cysts of histolytica in the faeces; thus, amongst the men they
were noted at the first and seventh; first,, second, eight and ninth;
second, third, fourth, fifth, sixth and ninth; second, fourth and
sixth; second and third only, second and sixth; and so on. Among
the women on the second only; the fourth only; the fifth and ninth;
the second, third and seventh; the second, fifth, sixth and seventh.
These patients were not being treated for the presence of these
cysts, for none of them showed any clinical symptoms attributable
thereto, and the mere presence of them does not imply amoebic
dysentery either present or past, and certainly does not warrant
specific treatment. The danger is more, of course, for others, for,
though causing no symptoms in the host, they may nevertheless
produce acute dysentery in another.
In this small series of one hundred and two cases it is seen that
a larger number of histolytica carriers was found amongst the men
than the women, in the proportion of three to two. Age appears to
have no significance; thus, of the females one was aged 13, two were
28, one eadh at 30, 50 and 56 years. Of the men the youngest was
23, then came one at 25, one each at 29, 30, 31 and 32, two at 35
and one at 52 years; the only point which may be mentioned is that
seven of the fifteen were found in patients during the third decade.
126
The importance of this infection with histolytica is great. From
an examination of a large number of cases, Dobell stated that the
incidence of infection does not appear to be conspicuously greater
among the cases arriving at the general hospitals for treatment than
among those reaching their final depots after such treatment. It
would seem, therefore, that the treatment which most of the dysenteric
patients had received had not been sufficient to rid them of their
infections. This refers to patients who suffer or have suffered from
an attack of dysentery. Seeing that the infection is high in my
series of patients who had not been abroad (i.e . 9 outside Jamaica)
and who showed no clinical symptoms of intestinal mischief, there is
every likelihood of greater spread now that large numbers of the
contingents sent to the war from that island have returned, and that
several of them suffered from dysentery in Egypt and elsewhere.
Further, 4 Egypt and its neighbouring lands are notorious centres of
amoebic infection. 1 Dobell has also shown that passage through ah
infective area, by affording an opportunity of exposure to infection
with histolytica , resulted in the acquisition of infection by a large
number of individuals. The incidence of infection in men returning
from Egypt and Gallipoli was not conspicuously higher among those
who were classified as ‘ dysenteries * than among those invalided for
other reasons. This is very applicable to the men who went from
Jamaica, for many (if not most) of them served in Egypt, and some,
at least, must have brought back infection with them.
As a matter of fact, the danger of spread is greater there than at
home, for many of those harbouring the cysts appear to be in perfect
health, are able to undertake their regular duties, associate freely
with their fellowmen, and may spread the disease. Also the duration
of carriage in a healthy carrier is not known, and would appear to be
almost indefinitely prolonged; though there are intervals when the
condition clears up, still this disappearance is only temporary. The
two main safeguards against spread are a good sewerage system and
abatement of the fly nuisance. Except in Kingston itself, and in
parts only of the city, there is no proper sewerage system, the street,
gutters constituting the sewer for many. As regards the fly question,
this is always a difficult problem in the tropics, and since we know
that flies can take up histolytica cysts, pass them unchanged through
their bodies and so infect food, the danger is clearly a very real one.
127
Wenyon and O’Connor’s (1917) remarks on the fly question may
be briefly quoted here: —
1. Flies readily take up encysted and other forms of protozoa
into their intestines.
2. Encysted forms remain in the intestines of the fly as long as
there is any faecal matter. If prevented from feeding they may
retain the cysts for as long as forty-two hours; if feeding is allowed,
they do not retain the cysts as long, and the flies may depbsit
material (and with it cysts) ingested only five minutes previously.
3. Cysts do not degenerate to any extent in the intestines of the
fly, but readily pass unaltered.
QiARDiA iN TBS TINA LIS
There is no proof that Giardia is pathogenic, at any rate in
adults. It has been found in a considerable number of instances in
this series, sometimes in a free state, sometimes encysted and in very
large numbers. Its presence has been noted in thirty-six of the
one hundred and two cases, or 35*29 per cent.; of these twenty-one
were amongst the fifty-three males and fifteen amongst the forty-nine
females. This, again, is a higher percentage than was found in
cases recorded by Dobell, Wenyon and O’Connor. The first-named
found 27*3 per cent % with a minimum of six examinations, but states
that this was probably not more than three-fourths of the real
number.
This parasite is usually found more frequently in patients
suffering from diarrhoea, but it is probably nearer the truth to say
that the diarrhoea or loose action is the ‘ cause * of the detection of
the infection than vice-versa , that the infection is the * cause ’ of the
diarrhoea. Two reasons may be offered to account for the high
percentage of infection in my series; firstly, that in order to ensure
the procuring of specimens many of the patients were given salines,
thus producing a loose action, and secondly, the concentration
method allowed of the discovery of the organism in cases where they
would otherwise have been so few as to be overlooked.
In reports of examinations carried out in England, it has usually
been found that Giardia was much more common in children, so
128
much so that Matthews and Malins Smith suggest that this flagellate
is mainly a parasite of children and that it may disappear from the
intestine in course of time.
In my series it was found in all cases examined (six) below the
age of 15 years, but was also present in others of 20, 56, and 60
years of age (five out of eight in the sixth decade); among the females
it was found in two of the three under 15 years of age, but also in
seven out of seventeen between 21 and 30 years, and in one each at
37 , 48 and 56 years.
CHiLOMASTIX M ESN I LI
Cysts of this parasite were found six times among the males and
seven among the females, giving a total percentage of 1274. In
some instances they were exceedingly numerous, occurring not only
in every field, but sometimes more than one in the field, the concen¬
tration method accounting in part for this. They are probably of
no pathogenic significance.
Lastly, it may be mentioned that Balantidium coli was found
twice: in a woman of 56 and a man of 22 years of age, in the latter
as the only parasite detected, in the former associated with E. coli.
The appended tables give succinctly the chief points in connection
with this investigation. The patients have been arranged in order
of age for ease of reference.
REFERENCES
Cropper, J. W., and Row, R. W. H. (1917). A method of concentrating Entamoeba cysts in
stools. Lancet , Vol. CXC, pp. 179-182.
Dobell, Clifford (1917). Amoebic Dysentery and the Protozoological Investigation of
Cases and Carriers. Med. Res. Committee , Special Report Series , No. 4.
Smith, A. Malins, and Matthews, J. R. (1917). Further Records of the Occurence of
Intestinal Protozoa in non-dysenteric cases. Ann. Trop. Med. & Parasitol Vol. IX,
pp. 183-193.
Wenyon, C. M n and O’Connor, F. W. (1917). Human Intestinal Protozoa in the Near
East.
Balantidium colu C.= Entamoeba coli. Cb.= Cbilomastix mesnili. G.= Giardia inuuinalis. H,= Entamoeba histolytica ... signifies that the stool was not obtained, —signifies negative examination.
I29
Table I —continued
130
Tabu II —Shewing rctulta of faecal examinations. Males* Jamaica.
Balantidium coli. C.=* Entamoeba c*li. Cb. =- Cbifamasti* mesnili. G.^Giardia intestinalis. H. = Bntamoeba histolytica. ... signifies that the uool was not obtained, —signifies negative examination .
29 Lymphadenitis
Table II — continued .
132
■33
A STUDY OF THE SIZES OF ENTAMOEBA
HISTOLYTICA CYSTS AMONGST
SYMPTOMLESS CARRIERS IN JAMAICA
BY
HENRY HAROLD SCOTT,
M.D., M.R.C.P. (LoncL), F.R.S.E., D.P.H.
GOVERNMENT BACTERIOLOGIST, HONG KONG } LATE GOVERNMENT BACTERIOLOGIST,
JAMAICA, B.W.I.
(Received for publication 13 April , 1921)
This investigation was carried out coincidently with another
which was undertaken to determine the incidence of intestinal
parasites amongst patients in hospital for some cause other than
dysentery or other intestinal complaints. Of the various patients
whose stools were examined, only those with a considerable number
of cysts will be dealt with in detail; others will receive briefer mention
but will be included in the total of all the cysts measured.
Malins Smith (1918) has shown that the curve obtained by
measurement of one thousand cysts of Entamoeba histolytica from
thirty cases is bimodal, between 7/* and 8/1 and between 12/* and
13/1; while that for Entamoeba coli is unimodal, between 16/* and
17/i. He also found that cases with cysts averaging between 9/*
and 10/* were very rare. He demonstrated that the two chief strains
were the ‘small’, averaging 7*7/*, and the ‘ordinary*, averaging
12*6 ft, and that in England infections with the former were rare in
those who had never been out of the country.
It was his experience also that each strain persisted for a long
time if left without treatment, or without being effectively treated,
and that the strains did not replace each other.
Dobell and Jepps (1918) claim to have established that there are
five strains of Entamoeba histolytica cysts, those occurring most
frequently being approximately 6*6/*, 8*3/*, 116/*, 13 3/* and 15/*.
‘For the complete demonstration of the existence of strains in
cysts of Entamoeba histolytica it is necessary to prove that the mean
diameter of cysts from any patient is not subject to any considerable
variation from day to day, but remains constant.*
*34
As regards the question as to whether change in size of cysts in
Entamoeba colt can occur from day to day, or rather from one day
to another, there is not sufficient evidence at hand, according to
Malins Smith.
In another paper by the same author (1919), he states that
samples of one hundred cysts (he is speaking of Entamoeba
histolytica ), taken on different days, often differ much more in
average size than samples from the same stool, so that these
differences cannot be accounted for by errors in sampling, and it has
been suggested by Dobell and Jepps ( loc . cit .) that emetine treat¬
ment may affect the size of the cysts, in that if they appear after
treatment they are liable to be larger than on normal occasions. In
the series dealt with below this question of the effect of emetine does
not come into discussion at all, since none of the cases showed any
intestinal symptoms to warrant interference, and they were not given
any treatment for the eradication of the cysts.
When mixtures of ‘ small ’ and ‘ ordinary * strains are found it is
only natural to expect that the cysts may appear in different propor¬
tions on different days, and thus there will be differences in the
average size of the cysts.
The criteria relied upon for differentiation of the cysts were those
stated in a previous paper on the incidence of intestinal parasites in
symptomless carriers in Jamaica, namely, the size and shape of the
cyst, the characters of the cyst-wall, the number of nuclei and
arrangement of the nuclear chromatin, the cytoplasm and its
inclusions—chromatoid bodies and vacuoles.
One of the most interesting of the cases in which cyst measure¬
ments were made regularly and in considerable numbers was that of
I. E. H., male. At his first examination cysts of both
Entamoeba colt and E. histolytica were found in fairly large
Fic. 1. Showing size of E. histolytica cysts (72). Case I, First examination.
7 8 9 10 11 12 13 14 15 16
135
numbers; one hundred and thirty-two coli and seventy-two
histolytica were measured. To save a long description, the curve
(following the method of Matthews and Malins Smith) below gives
at a glance the relative numbers present of cysts of the different sizes.
To avoid making the curve too complicated, the abscissae are
drawn at intervals representing I /*, and the ordinates at intervals
representing io units.
At the second examination, three days later, the numbers of cysts
found were less, but still considerable; the histolytica , however, were
rather more often met with, in the proportion of six to five, whereas
in the previous examination the coli were nearly twice as numerous
as the histolytica (this may, of course, be a mere sampling
coincidence). As regards the coli cysts, there is not much difference
to notice, except that those most frequently met with on the first
day were between i6/i and 17/1, constituting nearly 50 per cent,
(sixty-one out of one hundred and thirty-four), whereas on the next
occasion thirteen out of fifty were between 17/i and 18/1. As regards
the histolytica cysts, those most frequently met with, and those
nearest approaching them, were practically the same as on the first
occasion.
Fig. 2. Showing size of E. h istoly tic a cysts (60). Case I. Second examination.
This patient then left hospital for a week to see after some
private matters at home; further examinations were carried out on
his return. Not only were coli cysts quite infrequently met with,
but histolytica were numerous; in the course of measuring one
hundred of the latter, only twenty of the former were encountered.
More than one-third (seven) of the coli cysts were 19*4/* in diameter,
and the average size of all was i8'64/i; only one was found of the
size (16 fi to 17/i) which was most frequent at the first examination.
Also, as the curve below shows, the histolytica cysts were distinctly
smaller, though still nearer the size of the * ordinary * strain than the
‘ small \ Thus, at the first examination the most frequent were
136
12*6 fi and 13*2/*, while they were now at.the third examination
io*6/* and 11*2/*.
Fig. 3. Showing size of E. histolytica cysts (100). Case I. Third examination.
7 8 9 10 11 12 13
Two days later a fourth examination was made of the stools from
this patient, and again histolytica cysts were found in fair numbers,
while coli cysts were conspicuous by their absence. While measuring
two hundred of the former, only five of the latter were encountered.
Though among the histolytica cysts the size most frequently met
with was that of diameter 10*6/*, as before, nevertheless, as the
curve shows, those a little smaller still were present in considerably
greater proportion than on previous occasions. It is a matter of
much regret that this patient left hospital just afterwards, so that
I was not able to obtain further specimens, for though, as already
stated, the strains do not replace each other, nevertheless, these
Fio. 4. Showing size of E . histolytica cysts (200). Case I. Fourth examination.
7 8 9 10 11 12 13 14 15
137
findings are at variance with the statement that ‘ the mean diameter
of cysts from any patient is not subject to any considerable variation
from day to day, but remains constant.*
Thus, the sizes 12*6 p and 13*2 fi most frequent at the first
examination became at the third examination, a week or ten days
later, 10*6/1 and 1T2/i; and cysts 13*2/1 in size, the numbers of which
(viz., twenty-seven and fifteen) were only one less than the most
frequent numbers (viz., twenty-eight and sixteen) in the first and
second specimens examined, had dropped at the fourth examination
to only 1*5 per cent, of the cysts found.
Fig. 5> in which the curves for the different examinations are
placed on one chart, the proportions of cysts of various sizes being
reduced to percentages, shows well the change which occurred in the
size of the cysts in the course of a fortnight or so.
Fig. 5. Showing size of E. histolytica cysts. Case I. Four examinations.
• 8 9 10 11 12 18 14 15
- =3 first — - — - — =* second .. third = fourth
Fig. 6 gives the curve for the whole four hundred and thirty-two
histolytica cysts measured in this case.
Fig. 6. Showing size of E. histolytica cysts (432). Case I.
* 3 «
II. M. W., female. Several examinations were made also of
this case, and, except on one occasion, histolytica cysts were much
more numerous than coli cysts. On the first occasion on which
one hundred were measured,. 25 per cent, were of a diameter of
7‘9/«; this was the size most frequently met with, while the mean
for the one hundred was 8*94/*, and thirty-five were between 7 p and
8/1 in diameter.
Fig. 7 shows the curve of distribution on this occasion.
Fig. 7. Showing size of E. histolytica cysts (100). Case II. First examination.
8 9 10 11 12 13 14
On comparing these findings with subsequent ones, the condition
is most interesting, and I can find no other explanation than that
either this patient was infected with both the 1 small 9 and ‘ ordinary 9
varieties, or rather with the ‘ small * and one or more of the larger
varieties, and that after the first occasion the ‘ small * failed to
appear, or, secondly, that a change took place of the nature of
conversion of the small into the larger. This, it is generally held,
does not occur, though, as already stated, Dobell suggests that after
emetine treatment, if cysts reappear, they may be larger than before
treatment. In this case, as with the others, no emetine at all was
given.
There is one other explanation of the findings, namely, that the
wrong stool was sent up on the first day, as the findings then were
so different from those on subsequent occasions, but every care was
taken to avoid such a mistake. The patients were told what was
wanted and the reason for it, and the nurses were also asked to take
particular care to obtain the specimens fresh and send them up as
soon as possible after the passing of the stool.
Moreover, on comparing the figures for the different examina¬
tions, it will be seen that the transition of the mode of the curve is
gradual. Thus, in fig. 7 the greatest number had a diameter
139
between 7 fi and 8/1; after an interval of two examinations in which
no histolytica cysts were found at all, they appeared in large
numbers, those most in evidence having a diameter between 9 fi
and iOft.
Fig. 8. Showing size of E . histolytica cysts (100). Case II. Second examination.
8 9 10 11 12 18 14
On the third occasion those most frequently encountered had
an average diameter between 10 fi and lift, and in addition
there were more cysts between 13/1 and 14/* than at the previous
examinations.
Fig. 9. Showing size of E. histolytica cysts (125). Case II. Third examination.
8 9 10 11 12 13 14
At the fourth examination, at which the cysts were numerous,
those between 10/1 and 11 /1 were again the most frequent, and even
more so than at the previous examination.
Fig. 10. Showing size of E. histolytica cysts (130). Case II. Fourth examination.
8 9 10 11 12 13 14
140
The fifth examination showed a similar state of things as regards
those between 10 fi and 11/1, while those between 13/u and 14/1 were
becoming less again.
Fic. 11. Showing size of E. histolytica cysts (100). Case II. Fifth examination.
50
40
30
20
10
8 9 10 11 12 13 14
Finally, a short time before the patient left hospital the number
of cysts generally was less, but though those with diameter between
10 /i and 11 fi comprised about 25 per cent., there was a still
proportion, namely, fifteen out of forty (*.*., 37 per cent.) with a
diameter between 12 fi and 13/1, i.e. t the size known as ‘ordinary.*
We note, therefore, the peculiar fact that in the course of a month
there was a change from cysts of the 4 small * variety being most
numerous to those of the 4 ordinary * variety, passing through
intermediate phases.
This is graphically shown in fig. 12, which gives the superposed
curves of the six examinations, where the shifting of the mode to the
right is very evident. It will be noted that the curves for the third,
fourth and fifth counts strongly resemble one another. x
Fig. 12. Showing size of E. histolytica cysts. Case II. Six examinations.
50
40
30
20
10
8 9 10 11 12 13 14
= first second .= third ------- - = fourth
- -. — — = fifth = sixth
Fig. *3 gives the single curve for the whole five hundred and
ninety-five cysts measured from this case.
Fig. 13. Showing size of E. histolytica cysts (595). Case II.
8 9 10 11 12 13 14
III. C. H., male. This patient's stools showed cysts of
Entamoeba histolytica on two occasions during the three weeks, the
second being five days after the first. On the first, one hundred
cysts were measured, and, as the following curve shows (fig. 14),
those most frequently met with had a diameter of 112 /a, while
47 per cent, were between 11/1 and 12/1, and the mean for the
one hundred was 1132/1.
Fig. 14. Showing size of E. histolytica cysts (100). Case III. First examination.
8 9 10 11 12 13 14
On the other occasion on which histolytica cysts were found, out
of the fifty measured there were ten each of 9*3/1 and 11*5/1, and an
average diameter of 10*2/1; the greatest number (26 per cent.) were
between 9/1 and 10/1.
A curve of all the histolytica cysts measured from this case
(one hundred and fifty) is given in fig. 15.
H 2
Fic. 15. Showing size of E. histolytica cysts (150). Case III.
60
50
40
80
90
10
8 9 10 11 12 18 14
IV. T. M., male. On the first occasion on which cysts were
found in specimens from this patient, histolytica and Giardia only
were met with, and of the former very few—twenty only in a
prolonged search. Of these between one-third and a half showed
asymmetry, so that the mean of the two diameters was taken as
the measurement; of the twenty there were six each between 7 n and
8 ti and between 8/a and 9/u, the remaining eight being between
I I/a and I2/a.
For five days after this Giardia only was found, and then
histolytica was encountered in considerable numbers. One was able
to measure one hundred with very little searching, and during the
measuring of these a few coli cysts were also found, but only five of
them. The distribution of sizes of the one hundred histolytica cysts
is shown in fig. 16. As with the twenty seen on the first occasion,
only six were of the small size, the greatest number being between
11 H and i2/u as before, but this only exceeded by one that of those
between 10/1 and lift.
Fig. 16. Showing size of E . histolytica cysts (100). Case IV. First examination.
They were not found again till four days later, when practically
the same condition of things was noted and cysts were encountered
*43
in the same relative proportion, namely, six coli to one hundred
histolytica . Those met with most frequently were of the same size
as before, but the mean was higher, ir6ju as against io’8fi on the
previous occasion, a difference, possibly, not more than would be
accounted for by sampling errors with so small a number as
one hundred.
Fig. 17* Showing size of E. histolytica cysts (100). Case IV. Second examination.
8 9 10 11 12 13 14
The following five cases do not call for much individual or
detailed description. In all of them histolytica cysts were only
found occasionally, perhaps at one examination; their claim to
mention consists in the fact that when they were found one hundred
cysts were measured, so that a percentage of size was obtainable.
V. C. M., male. At the first examination a few coli cysts only
were found; during the next twelve days no cysts of any kind were
seen, then coli in small numbers. Giardia very numerous and
one hundred histolytica were measured. During the succeeding
twelve days again no cysts were seen in spite of repeated examina¬
tions, but then a few coli again made their appearance.
Fig. 18. Showing size of E. histolytica cysts (100). Case V.
8 9 10 11 13 18 14 16
VI. R. W., male. On two occasions only in three weeks were
cysts found in the stools of this patient. At the first there were not
/
i+4
many of either form, but twenty-six coli were found to twelve
histolytica . On the second occasion, eleven days later, almost the
same number of coli were seen, but histolytica were more numerous,
one hundred being measured. Those most frequently met with were
between 12 n and 13/1, with a mean for the one hundred cysts of
12*3 fi. This corresponds, therefore, to the 1 ordinary 9 type of cyst
infection.
8 9 10 11 12 18 14 15 16
VII. W. F., male. The occurrence of cysts in this patient was
very inegular. At almost the first examination histolytica cysts
were met with in considerable numbers, and one hundred were
measured. For the next four days none were seen at all; then a few,
eight only, were found after a long search. They were again absent,
or perhaps it were better to say undiscovered, for nine days, and
then in small numbers only and with Giardia cysts. Two days
afterwards coli cysts appeared for the first time since the examina¬
tion was begun, though not in large numbers, and Histolytica was
present in small numbers as before, Giardia being numerous. A
curve is given for the occasion on which one hundred cysts were
measured. Those most frequently met with had a diameter of
ir6/u, and the mean was 11*69^.
Fic. 20. Showing size of E. histolytica cysts (100). Case VI h
8 9 10 11 12 13 14 15
VIII. M. S., female. On four occasions histolytica cysts were
seen in this case; the interval between two appearances was ten days
in one instance. Three times out of the four a few cysts only were
seen, namely, nine, six, and fifteen, but once one hundred were
*45
found and measured. Coli cysts were usually present too, but not
in large numbers, and on one occasion none could be found at all.
Fig. 21. Showing size of E. histolytica cysts (ioo). Case VIII.
.9 10 11 13 13 14
IX. M.S., female. This patient left hospital after two examina¬
tions. At one no cysts were detected at all, at the second histolytica
cysts were found to be fairly numerous, while coli were present also
but in smaller numbers (one to four); nearly half of the histolytica
varied between lift and 1 2ft in diameter.
Fig. 22. Showing size of E, histolytica cysts (ioo). Case IX.
X. Finally, there may be mentioned a case in which histolytica
cysts were found on several occasions in conjunction with coli , but
at no time were they numerous. Fifty were measured twice, and
altogether a total of one hundred and sixty-seven. Those most
frequently met with had a diameter between and 8/i, and next to
this between lift and \2p.
Fig. 23. Showing size of E . histolytica cysts (167). Case X.
8 9 10 11 12 13 14 15
146
From three other cases between fifty and one hundred cysts were
measured, while in five more a smaller number still were found.
Some of these patients left hospital before the investigation could be
completed, or were transferred to other wards and the stools were
not sent up regularly. It would serve no useful purpose, therefore,
to discuss these in any detail; the total has been included in the
two thousand, three hundred and eighty-nine histolytica cysts
measured. This number includes cysts from eighteen patients, of
whom ten contributed over one hundred cysts each, three between
fifty and one hundred, and the remaining five a smaller number. It
may be repeated once again that none of these patients gave a history
indicative of previous attack of acute dysentery (this does not
preclude their having had such an attack) and none were
suffering from any intestinal trouble during the time they were under
investigation.
Fig. 24. Showing size of E. histolytica cysts (2,389).
If one considers fig. 24, the curve would appear to be a multi¬
modal one, with a mode between 7*5/11 and 8/1 (corresponding to the
H7
1 small * variety) with an average diameter of 7 7/1; a second between
10and 10*5 ji; a third between lip and 11 '$/*; a fourth about
1 2 p, and a fifth between 13/u and 13*5/*.
It is worth noting that there is among this series no mode at
I2'6 fi as has been found by Matthews and Malins Smith at home,
but there is a very marked one practically corresponding to the
1 r6 p variety of Dobell and Jepps, and also one corresponding to
the 13*3/1 variety (I forbear to call them 'strains*) of the same
author-.
8 9 10 11 12 13 14 15 16
When, however, the figure is drawn up giving the numbers found
within certain intervals of a complete 1/1, the curve then appears as a
bimodal one (fig. 25), though the second mode does not occur
148
between 12/1 and 13/*, as Malins Smith found, but between up
and I2p.
The interpretation of this I cannot give, for further examinations
for a prolonged period of a large number of cases would be necessary
before any dogmatic statement could be brought forward. Whether
the prolonged infection with the cysts of Entamoeba histolytica
leads to their becoming reduced in size, or whether the strain
in Jamaica is not the same as the ‘ordinary 1 strain found at
home, but a little smaller, is mere conjecture and not* worth serious
consideration with the small data at present available.
REFERENCES
Dobell, Clifford (1917). Amoebic Dysentery and the Protozoal Investigation of cases and
carriers. Med. Res. Committee , Special Report Series , No. 4.
-(1918). Parasitology. Vol. X, p. 320.
Matthews, J. R. (1918). Observations on the Cysts of the common Intestinal Protozoa of
Man. Ann. Trop. Med. & Parasit ., Vol. XII, No. 1, pp. 17-24.
Smith, A. Malins (1918). Measurements and Observations upon the Cysts of Entamoeba
histolytica and of Entamoeba coli. Ann . Trop. Med. & Parasit Vol. XII, pp. 27-69.
-(1919). A Contribution to the question of the number of Races in the species
Entamoeba histolytica. Ann. Trop. Med. & Par suit ., Vol. XIII, pp. 1-16.
■49
A STUDY OF THE SIZES OF ENTAMOEBA
COLI CYSTS AMONGST SYMPTOMLESS
CARRIERS IN JAMAICA
BY
HENRY HAROLD SCOTT,
M.D., M.R.C.P. (Lend.), F.R.S.E., D.P.H.
GOVERNMENT BACTERIOLOGIST, HONG KONG ; LATE GOVERNMENT BACTERIOLOGIST,
JAMAICA, B.W.I.
(Received for publication 13 April , 1921)
An investigation of the cysts of Entamoeba coli was undertaken
simultaneously with that of Entamoeba histolytica and on similar
lines. The stools were obtained from two series of patients, three
each of male and female, on alternate days, the patients being chosen
from those in hospital for some condition other than an intestinal
complaint. One series was sent up for examination on Mondays,
Wednesdays and Fridays, the other on Tuesdays, Thursdays and
Saturdays, for a period of three weeks.
It has been stated that, in contradistinction to the curve for
Entamoeba histolytica cysts, the curve for the cysts of Entamoeba
coli is unimodal, with the mode between 16 fi and 17^. Another
statement which has been made is that ‘the evidence upon the
important question as to whether change in cyst size [of Entamoeba
coli ] occurs from day to day is insufficient.’ The data founded on
the following series of cases, though not very extensive, may offer a
little evidence to help in the solution of this question.
As in the case of histolytica , each patient from whose faeces a
considerable number of cysts was measured will be mentioned
separately, and then a curve will be given for the whole number.
I. This patient exhibited both coli and histolytica cysts in
considerable numbers; the latter have been dealt with in a previous
paper. As regards the coli cysts, one hundred and thirty-four
were measured on the first occasion; among these the size most
frequently met with was 16*6/1 in diameter, and the mean for the
whole was 17*4/1. At none of the subsequent examinations, in spite
of the fairly abundant presence of histolytica % were coli cysts
plentiful; in fact, they became less and less frequent at each
examination. Thus, on the next occasion on which they were found
only fifty were measured, and a prolonged search resulted in finding
twenty and five respectively at the next two examinations.
Fig. i. Showing size of E. coli cysts (134). Case I.
15 16 17 18 19 90 91 99 93
15 16 17 18 19 90 91 99 93 94
II. The second case left hospital after two examinations had
been made, in both of which Giardia and coli cysts were fairly
numerous, particularly the former. On the first occasion one hundred
and fifteen coli cysts were measured, and on the second one hundred.
The curves for these and for the total are given below. At the
former examination those most frequently met with had a diameter
of 22*7/1, but these only exceeded those with a diameter of 19*8/1
by one, with a mean of 20*4/1; at the second, those of 19*8/1 were
most numerous, and the mean of the one hundred was 19*9/i.
Fig. 3. Showing size of E. coli cysts (115). Case II. First examination.
15 16 17 18 19 20 21 22 23 24 25 26 27
Fig. 4. Showing size of E. colt cysts (100). Case II. Second examination.
15 16 17 18 19 20 21 22 23 24 25 26 * 27
life At the first examination of faeces from this patient several
specimens and prolonged search revealed only seven histolytica and
three coli cysts. Two days later there was a different picture;
colt cysts were many, Giardia and Chilomastix were in considerable
numbers, while no histolytica were seen at all.
Again, after an interval of two days ho Giardia were found,
there were a few Chilomastix , three histolytica and four coli cysts
after long search. For the next week only Giardia was found, and
then coli were again numerous, as were also Chilomastix and
Giardia , and a few histolytica .
On the first occasion, where one hundred and two cysts were
measured, those most frequently met with had a diameter of 20*3/1,
the mean being 20*1/1; at the last examination 17 8/1 were the most
frequent, and the mean was 18*17/1.
Fig. 6. Showing size of E . coli cysts (102). Case III. First examination.
Fig. 7. Showing size of E . colt cysts (100). Case III. Second examination.
15 16 17 18 19 20 21 M 28 24 25 34
IV. The fourth patient's stools contained coli cysts on several
occasions, and in large numbers on some. On three of these
one hundred or more were measured, and the relative proportions of
the various sizes are shown in the accompanying curves. Those
occurring most frequently were 20*9/4, 19.8/4, and 19.8/4, respectively
with corresponding means of 19 8/1, 19*2/1, and 19*9/1, considerably
larger than the usual mean of 16/1 to 17/1 given for coli cysts in
general.
! 53
Fig. 9. Showing size of E . coli cysts (102). Case IV. First examination.
_1
17 18 1$ 20 21 22 28 24
Fig 10. Showing size of E . colt cysts (100). Case IV. Second examination.
14 15 16 17 18 19 20 21 22 23
Fig II. Showing size of E. coli cysts (100). Case IV. Third examination.
17 18 19 20 21 22 23 24
Fig. 12. Showing size of E. coli cysts (309). Case IV.
iS+
The following eleven cases exhibited Entamoeba colt cysts in
their faeces, but not sufficiently often for large numbers to be
measured on several occasions; a total of over one hundred was
measured from each, and, therefore, a brief remark on them is
warranted.
V. This patient has already been mentioned in the paper on
the measurement of cysts of histolytica , as she was passing both
histolytica and coli in considerable numbers, but, except on one
occasion, the former were in greater numbers.
At the first examination at which cysts were found, there was a
proportion of four histolytica to one coli . In measuring one hundred
of the former only twenty-two of the latter were encountered. Of
these the size most frequently met with (six in all) had a diameter
of about 21/1 (between 21*1/1 and 21*5/1; the average for the whole
was 20*1/1.
At the second examination prolonged search revealed only six
coli cysts, varying from 19*1/1 to 21*1/1 in diameter, with a mean
of 20*23/1. On this occasion prolonged and repeated search yielded
no histolytica cysts at all.
Two days later histolytica were again numerous while coli were
very scarce, only three being encountered during the time that
one hundred histolytica were measured. Yet again, after two more
days, though histolytica were fairly numerous, they did not exceed
the coli in such proportion, seventy-six of the latter being found to
one hundred and twenty-five of the former. On this occasion the
average size of the cysts was large, namely, 21*06/1, the most
commonly occurring being 22*4/1, of which there were eleven^ while
of 20*5/1 and 237/1 there were ten each. Two days later they
were present in about the same proportion, seventy-two coli to
one hundred and thirty histolytica ; some coli both larger and smaller
than on previous occasions were found, but otherwise the numbers
were fairly evenly distributed, the mean for the whole being 20*1/1,
the highest number being nineteen between 26/1 and 21/1.
At the next examination coli were present in a proportion of one
to three (thirty-three coli to one hundred histolytica ). There is
nothing particular to remark about them on this occasion; the most
commonly occurring had a diameter of 20*7/1, while only one less
were 20*1/1 one the one side and 22*5/1 on the other; the average for
the whole was 21*47/1.
X S5
At the last time of examination there were rather more coli than
histolytica (fifty-five to forty), the most common was 19*6/1, and the
mean of the whole was 19*79/1.
60
50
40
30
30
10
14 15 16 17 18 19 30 31 S3 33 34 35 36 37 38 39 30 81
VI. In this patient colt cysts in considerable number were found
on one occasion only, though histolytica were met with more
frequently. When they were in sufficient numbers one hundred were
measured, and the curve below gives the various sizes encountered.
It is a matter of regret that during this patient’s subsequent stay
in hospital these cysts were not again found, because it would have
been interesting to see whether this bimodal form of curve persisted,
in contrast with what usually obtains in infections with Entamoeba
coli cysts.
Fig. 14. Showing size of E. colt cysts (100). Case VI.
30
20
10
15 16 17 18 19 20 21 22 23
Fig. 13. Showing size of E. coli cysts (267). Case V.
r
y\
— 1
t
—
■
r
■
■
9
■
\
t
■
■
■
■
g
3
A
m
m
—
—
—
—
1
i
E:
—
■
9
9i
VII. This patient left hospital after one examination had been
made; at this, however, one hundred and eight coli cysts were
measured, of which the diameter of the most frequently .occurring
was I7’9M, with 17*5/1 only one less, and the mean was 17-8/1. The
curve of these is given (fig. 15).
Fig. 15. Showing size of E. coli cysts (108). Case VII.
16 17 18 19 20 21
156
VIII. On two occasions only were coli cysts found in any
numbers; on the first of these one hundred were measured, on the
second fifty. At subsequent examinations merely three or four were
found, and sometimes none at all. These two occasions were but
two days apart, and it is interesting to note the difference in the
prevailing size. Thus, with the first one hundred the diameter of
those most frequently met with was 19*2/1 and the mean was 19*75/*,
while at the second time of examining the commonest was 15*9/* and
the mean 17*83/*. These wide differences illustrate remarkably the
errors possible from the measurement of small numbers of cysts. The
curve for the whole one hundred and fifty is given here (fig. 16).
«
Fig. 16. Showing size of E . coli cysts (150). Case VIII.
IX. After several negative examinations coli cysts were found in
this patient’s stool on one occasion when one hundred were measured.
The most common had a diameter of 17*8/* with 19 8/* only one
less, and the mean was 18*4/*.
Fig. 17. Showing size of E. coli cysts (100). Case IX.
X. This patient unfortunately left hospital just after the first
positive examination. Coli cysts were fairly numerous, one hundred
being measured, but the noteworthy point is the tendency for
the majority to be of a somewhat small size, as shown by the
accompanying curve, though the mean for the whole works out at
i6‘2 fi. It is a matter for regret that I was not able to follow up
this patient and obtain further specimens.
Fig. i 8 . Showing size of E . colt cysts (ioo). Case X.
80
20
10
14 15 16 17 18 19 20
XI. Cysts were never very numerous in this case and were
present intermittently, at least were only discovered intermittently.
One hundred and ten cysts were measured, of which the curve is
given (fig. 19).
A
L_
t
\
14 15 16 17 18 19 20 21 22
XII. Coli cysts were found on nearly every occasion in this case,
but not in large numbers; sometimes a bare half-dozen or so after
prolonged search, but on three occasions fifty were measured. On
the first and second of these the diameter of cyst most frequently
met with was 19*8/*, but on the third only 17*6/1. The number
investigated was not sufficient to enable one to say whether this was
a mere accident or error in sampling, but the smaller size was
encountered on the last occasion on which they were found prior to
the patient’s departure from hospital.
It is a matter of conjecture as to whether immature cysts were
being expelled before their final disappearance, or whether, as a
corollary of Dobell’s statement—that emetine-resistant histolytica
cysts, or cysts appearing after a course of emetine, are inclined to be
larger than before—when the patient is getting the better of
his infection, the cysts may diminish in size* One must not, of
course, lose sight of the fact that histolytica infections are pathogenic
whereas coli probably ard not. It may also be remarked, however,
that in this patient some of the largest met with in the investigation
were encountered during the last three or four examinations ; while the
largest during the first ten days* examinations was 24*4/1, towards
the last eight days there were seen a few as large as 26*3/1, 27*1/4
and 28*3/1, but with only eight nuclei. Fig. 20 gives the curve for
the total of one hundred and ninety-five cysts measured in this case.
Fig. 20. Showing size of E. coli cysts (195). Case XII.
SO
40
SO
20
10
15 16 17 18 19 20 21 22 28 24 25 26 27 28 29
XIII. In the stools from this patient cysts were scarce, but were
measured from time to time when seen, a total of one hundred and
nineteen cysts being obtained. The curve from this case is given
(fig. 21). The most frequent was 19*6/4, and the mean 201/4.
XIV. From this patient only sixty coli cysts were measured on
two occasions. The size most frequently met with had a diameter of
17*2/4, and the mean was 18*1/4.
XV. Cysts were found but once only in this patient’s stools and
l S 9
fifty were measured; twenty of them had a diameter of 17*6/1, and
the mean of the total worked out at 17*73/1.
In all, two thousand five hundred Entamoeba coli cysts have
been measured, and the charts below show the various sizes met with.
From these curves it will be noticed that instead of a unimodal curve
with mode between 16/1 and 17/1, it is distinctly bimodal, with
modes between 17/1 and 18/1 and between 19/1 and 20/1.
Figs. 1, 6, and 20 show the unimodal type; fig. 2 shows in
addition a smaller increase again between 19/1 and 20/1; the
majority of the rest show a bimodal graph, as stated above (figs. 5,
11, 19, for example); when the tendency is to be unimodal the mode
is between 19/1 and 20/1 (figs. 12, 16, 20); in figs. 8 and 13 the
majorities are between 17/1 and 18/1 and 20/1 and 21/1, while fig. 17
is unimodal between 17/1 and 18/1; fig. 21 shows yet another form.
No inference of value can be drawn from the individual curves
of these cases, for the numbers in each are too small, but the graph
of the whole two thousand five hundred (fig. 23) brings out the point
clearly that, judging from the present findings, the graph of the
sizes qf Entamoeba coli cysts is also bimodal, both modes being at
a higher level than has been recorded at home, namely 17/* to 18/*
and 19 fi to 20/*, instead of between 16/* and 17/*.
Fig. 22. Showing size of E. coli cysts (2,500).
560
Fig. 23. Showing size of E . coli cysts (2,500).
i6i
CESTODES FROM INDIAN POULTRY
BY
T. SOUTHWELL
(Received for publication 3 June , 1921)
With the exception of three species, all the Cestoda described below
were collected by Lt.-Colonel Clayton Lane, I.M.S. (retired), from fowls,
at Berhampore, Bengal, India, during 1912 and 1913. The worms were
deposited by the writer in the Indian Museum and were sent for identi¬
fication to the School of Tropical Medicine by Dr. Nelson Annandale,
Director of the Zoological Survey of India. *
The following is a complete list of Cestoda recorded up to the present
in fowls from all parts of the world :—
Metroliasthes lucida, Ransom, 1900 ?
Hymenolepis carioca (Magalhaes, 1898), Ransom, 1902.
Choanotaenia infundibulum (Bloch, 1779), Cohn, 1899.
Bothriotaenia longicoUis (Molin, 1858), Railliet, 1892 (recorded
. once only).
Cotugnia digonopora (Pasquale, 1890), Diamare, 1893.
Dicranotaenia sphenoides (Railliet, 1892), Railliet, 1896.
= Dicranotaenia cuneata (Linstow, 1872), Railliet, 1893.
Drepanidotaenia infundibuliformis (Goeze, 1782), Railliet, 1893.
Hymenolepis cantaniana (Pol., i860). Ransom, 1909.
= Davainea oligophora , Magalhaes, 1898.
Hymenolepis villosa (Bloch, 1782), Wolffh., 1899.
Davainea cesticillus (Molin, 1858), R. Blanchard, 1891.
cohni , Baczy, 1914.
„ echinobotkrida (M6gnin, 1880), R. Blanchard, 1891.
„ exilis (Duj., 1845), R. Blanchard, 1899.
„ longicoUis (Molin, 1858), Fiihrmann, 1908.
„ mutabUis , Reuther, 1901.
„ paraechinobothrida, Magalhaes, 1898.
= D. echinobotkrida ?
Davainea penetrans , Baczy, 1914.
„ proglottina (Davaine, i860), R. Blanchard, 1891.
= D. dubinis , Meggitt, 1916 ?
„ proglottina dublanensis (Kowl., 1894), Fuhrmann, 1905.
„ tetragona (Molin, 1858), R. Blanchard, 1891.
„ urogalli (Modeer, 1790), R. Blanchard, 1891.
„ varians , Sweet, 1910.
„ vigintivasus , Skriab., 1914.
„ volzi, Fuhrmann, 1905.
Fimbriaria fasciolaris (Pallas, 1781), Wolffh., 1899.
= F. malleus (Goeze, 1782), Froel., 1802.
= F. mitra f Froel., 1802.
Fuhrmann (1920) has recently split up the old genus Davainea
(Blanchard) into four genera, viz. :—
Davainea , Blanchard. Type D. proglottina , Davaine.
Davainoides , Fuhrmann. Type D. vigintivasus , Sk., and D.
polycalceola , Jan.
Houttuynia , Fuhrmann. Type D. struthionis (Houtt.).
Raillietina , n.nom.
The latter genus he splits up into four sub-genera, as under :—
ParonieUa , n.s.g. Type D. longispina .
Ransomia, n.s.g. Type D. tetragona (Molin).
Skrjabinia , n.s.g. Type Z>. cesticiUus (Molin).
Johnstonia , n.s.g. Type D. echinobothrida (M£g.).
Fuhrmann removes his genus Ophryocotyloides from the sub-family
Davaininae and places it in the sub-family Ophryocotylinae, Fuhrmann.
Meggitt (1921), in his paper on tapeworms from the ostrich, gives a
key to all the known species of Davainea .
The collection dealt with in this paper comprises the following
species:—
Metroliasthes lucida , Ransom, 1900.
Cotugnia digonopora (Pasquale, 1890), Diamare, 1893.
Dicranotaenia sphenoides (Railliet, 1892), Railliet, 1896.
Davainea cesticillus (Molin, 1858), Blanchard, 1891.
„ echinobothrida (M£gnin, 1880), Blanchard, 1891,
„ tetragona (Molin, 1858), Blanchard, 1891.
Monopylidium gaUinarum, n.sp.
Diorchis , sp. (americana? Ransom, 1909).
*63
Metroliasthes lucida , Ransom, 1900
Numerous specimens from a doipestic fowl, Gallus gall us domesticus ,
Angul, Orissa, India: collected by the author in 1912. Numbered
2 E V in the collections of the Indian Museum.
7
There appears to be some doubt whether this species has been
previously obtained from fowls or not.
Cotugnia digonopora (Pasquale, 1890), Diamare, 1893
1. Four specimens from a domestic fowl, Berhampore, Bengal,
collected by Lt.-Colonel Clayton Lane, I.M.S.
2. Two specimens as above ; collected on 29.5.13.
3. Three specimens as above, 1912.
Dicranotaenia sphenoides (Railliet, 1892), Railliet, 1896
= D. cuneata (von Linstow, 1872), Railliet, 1893
Five specimens from a domestic fowl, Berhampore, Bengal, collected
by Lt.-Colonel Clayton.Lane, I.M.S., in November, 1912.
Davainea cesticillus (Molin, 1858), R. Blanchard, 1891
1. Five specimens from a domestic fowl, Berhampore, Bengal,
collected by Lt.-Colonel Clayton Lane, I.M.S., 1912.
2. One specimen from Gallus sonnerata (Zoological Gardens, Calcutta)
collected by the author on March 3rd, 1917.
Davainea echinobothrida (M^gnin, 1880), R. Blanchard, 1891
1. Three specimens from two domestic fowls, Berhampore, Bengal,
collected by Lt.-Colonel Clayton Lane, I.M.S., 1912.
2. Numerous specimens (about 50) from a jungle fowl (GaUus
bankiva ), Berhampore, Bengal, collected by Lt.-Colonel Clayton Lane,
I.M.S. Numbered Z.E.V. in the collections of the Indian Museum.
7
5. Three specimens from Gallus ferrigincus, Zoological Gardens,
Calcutta, collected by the author on May 27th, 1915.
Davainea tetragona (Molin, 1858), R. Blanchard, 1891
1. A total of twenty-four specimens were collected from fowls on
seven different occasions, in Berhampore, Bengal, by Lt.-Colonel Clayton
Lane, I.M.S., in 1912.
The two preceding species are very closely related and Stiles considers
them to be identical. T. tetragona , however, does not, it is stated, produce
164
the pathological effects which result from the presence of T. echinobothrida,
viz., nodular disease. .
Ransom, however, points out that D. echinobothrida is a larger form
than D. tetragona, possessing a larger head and rostellum, and larger
suckers, hooks, and cirrus pouch; the hooks are also more numerous in
D. echinobothrida than in D. tetragona . In mature forms these characters
appear to us sufficient to distinguish between the species.
Meggitt (1921) states that T . tetragona (Mol.) possesses ' unilateral
genital pores and D. echinobothrida (M^g.) alternating pores in the adult
form and unilateral pores in the young. The two species are closely related
and no classification can be regarded as satisfactory which separates
them. A character upon which a classification may perhaps be finally
based is the behaviour of the uterus and the origin of egg capsules/
Fiihrmann (1920) considers them distinct, his opinion being based
on the pores being unilateral in one and irregularly alternate in the other.
In this opinion we concur.
The following worms have not hitherto been described from fowls :—
Monopylidium gallinarum, n.sp.
Two specimens from a domestic fowl, Berhampore, Bengal, India,
collected by Lt.-Colonel Clayton Lane, I.M.S., in 1912.
External Anatomy. The worms measured about 21 mm. long,
and the greatest breadth was 2-5 mm. All the segments were much broader
than long, even the posterior ones being shallow. The number of segments
was about 130.
The head. The head is prominent and measures about 300 fi long
and 500/4 broad ; in both specimens the rostellum was retracted and
appeared as a small, bluntly-pointed projection, armed with about 120
hooks, each measuring about 90/4, and arranged in 2 rows. The suckers
are large, conspicuous, and unarmed. There is no neck.
Internal Anatomy (fig. 1). The outer longitudinal muscular
layer consists of a large number of separate dorsal and ventral strands.
The lateral water vessels are large and clearly seen, both in the entire
worm and in sections.
Genitalia. As only two worms were available, it was found impossible
to do more than work out the gross details of the anatomy.
Male genitalia. There are about 30 globular testes, each measuring
about 35/i, and these lie for the most part on each side of the ovary,
although a few lie in front of it.
F&s deferens. The cirrus pouch is large and muscular, and the genital
atrium runs anteriorly.
Female genitalia. The ovary is situated in the middle of the segment,
slightly posterior, and median to, the receptaculum seminis. It measures
about 225/4 broad and 60/4 long.
FicS. 1. Monopylidium gallinarum, n.sp., horizontal section. Ant., anterior } e.x., excretory
vessel; 0., ovary ; Pst., posterior ; r.i., receptaculum seminis j t. } testes ; v ., vagina. X 50.
Vagina . The receptaculum seminis measures about 45/4 by 15/4, and
lies on the pore side of the ovary.
The genital pores are unilateral and occur at the extreme anterior
corner of the segment. The eggs lie in capsules, each capsule containing
from five to nine. The outer egg-envelope measures about 35 fi and the
oncospheres about 25/ 4 .
Diagnosis. The number and size of the hooks, the position of the
genital pore, the fact that all the segments are much broader than long,
and the occurrence of numerous eggs in each capsule, differentiate this
worm from any other species of the genus Monopylidium .
Johnston (1911) discusses the relationship between the genera
Monopylidium and Choanotaenia .
We consider our species to be new and have therefore named it
M . gallinarum.
Diorchis sp. (americana ? Ransom, 1909)
Several specimens from a hen, Calcutta, Bengal, India, collected by
the author in 1915.
The only points in which our specimens differed from D . americana ,
Ransom, were as follows :—
Length
Breadth
Rostellar
1 hooks
Hooks on
suckers
Our specimens. ...
16 mm.
0*4 mm.
i absent
1
absent
D. americana .
20-25 mm.
o*6 mm.
Ten present !
present
i66
Two heads only were found amongst our specimens, and in both
cases the rostellar hooks were absent, although fragments were to be seen.
In spite of careful examination, no hooks were seen on the suckers ;
possibly they had worn off.
The uterus contained many eggs, but the oncospheres were not fully
formed. The outer egg-envelope measured about 25 fi and the embryo 16 fi.
No representative of this genus has previously been recorded from
fowls, although they occur occasionally in ducks, coots, sandpipers,
flamingoes, etc.
The opportunity is here taken of recording the following Echinorhyn -
chid , also obtained from an Indian Jungle fowl:—
Heteroplus grandis (Van Cleave, 1916), Van Cleave, 1918
= Mediorhynchus grandis , Van Cleave, 1916
Two females only from the intestine of Gallus sonnerati (South Indian
Jungle fowl), Zoological Gardens, Calcutta, collected by the author on
April 17th, 1917.
The type species was described from Quiscalus quiscula (the purple
grackle) and has since been recorded by the same writer from SturneUa
magna (the meadow lark) and Corvus brachyrhynchus (the crow).
Our specimens agree in detail with Van Cleave's account of this species
except in size. The females described by Van Cleave measured 27 mm.
to 35 mm. long, and the greatest breadth was 1*4 mm. Both our specimens
measure 80 mm. long, and have a maximum breadth of 3*5 mm.
REFERENCES
Baylis, H. A. (1918). A collection of Entozoa chiefly from birds from the Murman coast.
Ann. & Mag. Nat. Hist., Series 9, Vol. III.
Fuhrmann, O. (1908). Nouveaux Tanias D’Oiseaux. Revue Suisse de Zoologie , Vol. XVI.
- (1908). Die Cestoden der Vogel. Zool. Jabrb., Suppl. Vol. X, No. 1.
- (1913). Nordische Vogelcestoden aus dem Museum von Goteborg.
- (1918). Cestodes d’oiseaux de la Nouvelle Caledonie et des lies Loyalty. F. Sarasin
& J. Roux, Nova Caledonia. Zoologie , Vol. II, L, IV, No. 14.
- (1920). Considerations Generates sur les Davainea. Festschrift fur Zschokke, No. 27.
Johnston, T. H. (1909). On the anatomy of Monopylidium passerinum, Fiihrmann. Proc.
Royal Soc. of N.S. Wales , Vol. XLIII.
- (1911). New species of Avian Cestodes. Proc. Linn . Soc. N.S. Wales, Vol. XXXVI,
Pt. I.
Mrggitt, F. J. (1920). A contribution to our knowledge of the tapeworms of poultry.
Parasitology, Vol. XII, No. 3.
- (1921). On two new tapeworms from the Ostrich, with a key to the species of
Davainea. Parasitology, Vol. 13, No. 1.
Ransom, B. H. (1905). The tapeworms of American chickens and turkeys. U.S. Dept. Agric.,
Bureau of Animal Industry , Circular 85.
- (*909)- The Taenoid Cestodes of North American birds. U.S. Nat. Hist. Mus. y
Bull. 69.
Stiles, C. W. and Hassall, A. (1896). Tapeworms of poultry. U.S. Dept. Animal Industry,
Bull . 12.
167
CESTODES FROM AFRICAN RATS
BY
T. SOUTHWELL
(Received for publication 3 June , 1921)
1. Davainea celebensis, Janicki, 1902
Four specimens and some fragments were collected from a rat,
Mus rattus , by Dr. J. W. S. Macfie, at Accra, March 26th, 1921.
This species was first obtained from Mus meyeri in the Celebes, and
Janicki, in his account of its anatomy, says nothing about the head. We
therefore give the following description :—
The head measures about 150/4 long and 650/4 broad. The suckers
are abbut 170/4 in diameter and are unarmed. The rostellum is small,
measuring about 7Q/4 in diameter, and is armed with a single row of
60 hammer-shaped hooks measuring about 12/t.
The neck is about 3 mm. long.
Fiihrmann has recently split up the genus Davainea into four genera,
one of which (i.e., RaiUietina , nom.nov.) he has further divided into four
sub-genera. Of these sub-genera, our species falls under Ransomia , and
the worm is accordingly named Ransomia celebensis (Janicki).
2. Zschokkeella guineensis (Graham), 1909
= Thysanosoma gambianum, Beddard, 1911
— Thysanotaenia gambiana, Beddard, 1911
= Zschokkeella gambiana, Beddard, 1912.
Graham, in the ' Report on Plague in the Gold Coast in 1908/ was
apparently the first to refer to this species ; he named it Davainea
(Guineensis, sp.nov.) [sic].
Beddard, describing in 1911 the same worm from the same host, and
also from the same area, was apparently unaware of Graham's paper and
gave it the name Thysanosoma gambiana ; later in the year he changed
the name to Thysanotaenia gambiana, and in 1912 to Zschokkeella gambiana.
According to the rules of nomenclature this worm now becomes Zschokkeella
guineensis (Graham), Beddard, 1912.
i68
In our collection we have about 600 specimens from the following
sources:—
(1) From pouched rat, Cricetomys gambianus, Accra, December, 1914,
and February, 1915. Collected by Dr. J. W. S. Macfie.
(2) From Mus rattus and Mas decutnanus, Accra. No date. Col
lected by Dr. J. W. S. Macfie.
REFERENCES
FUhrmann, O. (1920). Considerations gtn£rales sur les Davainea. Festschrift fur Zscbokke ,
No. 27.
Graham, W. M. (1908). Ceatodes. Report on Plague in Gold Coast in 1908 by W. J. Simpson.
Janicki, C. V. (1920). Uber zwei neue Arten des genus Davainea. Arch. Parasitol ., Vol. VI.
Miggitt, F. J. (1921). On two new tapeworms from the Ostrich, with a key to the species of
Davainea . Parasitologyy Vol. XIII, No. 1.
A NEW SPECIES OF CESTODA FROM
A CORMORANT
BY
T. SOUTHWELL
(Received for publication 3 June, 1921)
Dilepis kempi, n.sp.
Four specimens from the little cormorant, Phalacrocorax pygmaeus
(= P. javanicus), North Lohtak Lake, Manipur, Assam, 14.2.20. Station I,
Manipur Survey. Collected by Dr. S. W. Kemp, Zoological Survey of
India.
External Anatomy. The largest worm measures 5 cms. long, and
the greatest breadth is about 1 mm. It contains over 500 segments,
all of which are broader than long ; the posterior segments measure about
900/i, broad and 400/4 long.
The genital pores are unilateral and situated in the anterior half of
the segment.
Head. The head is about 220/4 long and 400/4 broad. The rostellum
is about 170/4 long and about 160/4 broad ; it is armed with twenty hooks
arranged in two rows. The posterior row of hooks curve backwards
strongly and measure about 135/4; the anterior row are not so strongly
curved and measure about 175/4. The diameter of the suckers is about
IOO/4.
Neck. There is no neck.
Internal Anatomy. Male genitalia. There are three testes situated
in the median field. They measure about 140/4 by 70/4, their long diameter
being dorso-ventral. The vas deferens arises somewhat ventral and,
curving dorsally, runs in a lateral direction, dorsal to the water vessel and
ventral to the vagina, to the pore (figs. 1 and 2). The cirrus pouch is
small and insignificant.
170
Female genitalia . The ovary is situated in the median anterior field.
It consists of two irregularly-shaped wings, each measuring about 160^.
broad.
Fig. 1. Dilepis kempt , n.sp., transverse section, ex.d ., excretory vessel, dorsal; ex., excretory
vessel, ventral; l.m longitudinal muscles ; 0., ovary ; r.i., receptaculum seminis ; testes ; t.m
transverse muscles ; v. 9 vagina ; v.d., vas deferens ; v.g vitelline glands, x 100.
The vagina is conspicuous, running dorsal to the vas deferens and
water vessel. It opens in front of the vas deferens. It dilates a little into
a seminal receptacle (figs, i and 2), close to the ovary.
The vitelline gland measures about 70/4, by 50/i and lies posteriorly
between the two wings of the ovary (figs. 1 and 2).
Uterus. This is a large sac-like organ with very large and numerous
outpocketings, extending lateral to the water vessels on both sides. No
ripe eggs were seen.
Diagnosis. This species is distinguished from all other species of
Dilepis on account of the large size of the rostellar hooks and the occurrence
of three testes only. The presence of a double row of hooks shows that
the species does not belong to the genus Hymcnolepis.
We have named this species in honour of Dr. S. W. Kemp, Superinten¬
dent of the Zoological Survey of India, who collected the specimens.
REFERENCES
Fuhxmann, O. (1901). Die Cestoden der Vogel det weis*en Nils. Results o f the Swedish Zool .
Exp. to Egypt and the White Nile, 1901.
- (1908V Non re tux Tririas d’Oiseaux. Revue Suisse de Zoologie, Vol. XVI, No. 1.
-(1908). Cestoden der Vdgel. Zool . Jabrh., SuppL Vol. X, No. 1.
Johnston, T. H. (1912). New species of Cestodcs from Australian Birds. Mem . Queensland
Museum, VoL I.
Luhx, M. (1910). Susswasserfauna Deutschland*. Part 18 : Cestodea.
Ransom, H. B. (1909). The Taenoid Cestodes of North American Birds. US. National
Museum, Bull. 69.
I7J
A CASE OF SUSPECTED LEPROSY
BY
J. W. W. STEPHENS
AND
S. ADLER
(Received for publication 22 June, 1921)
Plate XVI
X.Y., age 16. Bom in England, 1904. Has resided in Capetown,
S. Africa, from 1905 until September, 1920.
History of Illness :—
April, 1920. Wasting noticed on dorsal side of left hand between
1st and 2nd metacarpals, and a stinging sensation in the little
finger.
June, 1920. Three spots noticed on left side of face and brownish
patches on both eyebrows. Later brownish patches noticed on
left forearm and hand, and on both lower limbs.
August, 1920. Noticed deformity and impairment of movement in
left hand.
Examination of the patient :—
1. A number of dry, scaly, brownish, thickened areas in part cir¬
cumscribed up to three-eights of an inch in diameter, in part somewhat
diffuse. None of these areas was anaesthetic except for those involved
in the areas of anaesthesia, to be described later. The distribution of
these is shown approximately in figs. 1 and 2.
2. A condition of claw hand (left). The actual condition can be
appreciated from the photographs (Plate XVI).
3. Thickening of the left ulnar nerve which in the ulnar groove
was about three times as thick as on the right side. The left internal
cutaneous nerve was palpable and about a quarter of an inch thick.
4. The left forearm and hand Were anaesthetic over an area shown
approximately in figs. 1 and 2.
5. The left hand was colder to the touch than the right.
From 30.12.20 to 14.4.21 during his stay in the ward there was no
rise of temperature.
27.1.21. The Wasserman reaction was completely negative.
On 7.2.21 and 21.2.21 the patient burned his left hand on the hot
water-pipes of the ward without being aware of it.
Examination for Leprosy Bacilli :—
1. One of the affected areas on the left forearm was excised at the
London School of Tropical Medicine, November, 1920, and examined by
Dr. Low for leprosy bacilli with negative result.
2. Numerous scrapings and punctures of the various skin lesions,
e.g., on the eyebrows, face, and of the nasal mucosa, were negative.
3. 2 c.c. of blood from a vein were haemolysed and centrifuged;
the deposit was examined for leprosy bacilli with negative result.
4. A piece of the internal cutaneous nerve was excised, and sections
and smears were stained for leprosy bacilli with negative result.
The excised nerve was examined for us histologically by Professor E.
Glynn, who reported that * there are three caseous areas apparently
corresponding to large nerve bundles, surrounded by fibrous tissue in
process of formation. No nerve fibres were seen. The appearance is
quite consistent with fibro-caseous leprosy.'
On re-examination of the patient on 14.6.21 the only change noticed
was a disappearance of affected areas on the dorsum of the hand and a
diminution in extent of some of the areas on the face.
In view of the fact that no evidence of the presence of leprosy bacilli
was obtained no treatment was adopted.
176
EXPLANATION OF PLATE XVI
Fig. 1. Ventral aspect of right and left hands. The left hand shows
(a) its smaller size; (A) ill-defined cutaneons furrows;
(c) wasting of the thenar and hypothenar eminences—flatness
of the hand ; (d) flexion of the interphalangeal joints of the
2nd, 3rd, 4th, and 5th fingers ; (e) indication of the thickened
rough skin areas at the base of 3rd, 4th, and 5th fingers;
(/) the scar of a bum at the base of the hand on the ulnar
side.
Fig. 2. Lateral view of the right and left hands. The left hands shows
(a) wasting hi the interspace between the 1st and 2nd meta¬
carpal bones ; (b) extension of the metacarpo-phalangeal
joints and flexion at the interphalangeal joints; (c) wasting
of the forearm.
Annals T:op. Med. & ParasitolVol. XI
C. Tin ling <5r* Co., Ltd., Imp
177
OBSERVATIONS ON THE
CERATOPOGONINE MIDGES OF THE
GOLD COAST WITH DESCRIPTIONS OF
NEW SPECIES .
PART IV
BY
HENRY F. CARTER
A. INGRAM
AND
J. W. S. MACFIE
(Received for publication J April , 1921)
Genus DASYHELEA , Kieff.
Ceratopogon y Wtz. {proparte) {tier. Mg. Edws.). Linnea Eniomologicae. Vol. VI,
1852.
Culicoidesy Kieff. et auct. {pro parte) (1 nee . Latr.).
Dasyhelea , Kieff. Bull. Soc. Hist. Nat. Med. Vol. Ill, 1911.
Prokempia , Kieff. Rec. Jnd. Mus. Vol. IX, 1913.
PseudoculicoideSy Mall. Bull. III. Sta . Lab. Nat. Hist. Vol. X, 1915.
Dasyheliay Brunetti. Rec. Ind. Mus. Vol. XVII, 1920.
This genus is closely related to Culicoides t Latr. It was erected
by KiefFer in 1911 for the Indian species D. halophila y and
subsequently (1913) divided by him into three sub-genera—
Dasyhelea (sens, stric.), Prokempia and Kempia —according to the
nature of the wing hairs and the empodium. Later (1917) the last
two sub-genera were given generic rank by this author, who then
associated Prokempia with Dasyhelea but removed Kempia to the
Atrichopogon group. Still later (1919} KiefFer again placed
Prokempia as a sub-genus of Dasyhelea. Prokempia apparently
differs from Dasyhelea only in the absence or relative scarcity of the
longer wing-hairs, and was separated for this reason; but although
the density of arrangement of these hairs is much more uniform in
Dasyhelea than in CulicoideSy this character cannot, in our opinion,
be considered of more than specific value.
i 7 8
Malloch, in 1915, founded the genus Pseudocultcoides for
Ceratopogon mutabilis , Coq., especially distinguishing it from
Culicoides by the absence of thoracic cavities and the antennal
structure of the male. The description and figures of this species
given by Malloch indicate that it is a typical Dasyhelea , and Kieffer
(1919) accordingly placed Pseudoculicoides as a synonym of the
latter. Malloch, however, made no reference to the eyes—the
hairiness of which was given by Kieffer as one of the diagnostic
characters of the genus—either in his generic definition or specific
description; but Mr. F. W. Edwards, in a private communication,
informs us that the eyes of specimens (named by Malloch) of
C. mutabilis in the British Museum collection are shortly but
distinctly hairy, and therefore Kieffer’s decision in regard to this
question must be accepted.
External Morphology.
The detailed anatomy of Dasyhelea differs from that of
Culicoides (see Part II of this study, 1920) as follows: —
Adults. Head. Eyes in both sexes densely clothed with
microscopic hairs.
Mouth-parts (fig. 1). Proboscis somewhat shorter than that
" d
Fio. I. Mouth part* of D. paUidihdur ($) : a —labmm ; b —maxilla ;
c —mandible; d —hypopharynx. ( x 490 circa.)
of Culicoides, the component organs, except the laEium, differently
formed and apparently not adapted for piercing in either sex. The
labrum is strongly* chitinised, broad at the base gradually tapering
179
to a rounded^ hairy apex. The hypopharynx is similar in shape to
the labrum, but tapers more rapidly and ends in a pointed apex;
the distal fourth bears numerous hair-like marginal processes. The
mandibles and maxillae are poorly developed and devoid of teeth;
the former are closely applied to (and not easily separated from) the
labrum and hypopharynx, and are enclosed in membranous sheaths
which extend considerably beyond their extremities; the maxillae are
rudimentary and are reduced to small, thinly chitinised, blade-like
structures. The palpi in both sexes appear to be composed of four
segments owing to the first being very rudimentary and often almost
indistinguishable; the third segment is usually distinctly longer than
the fourth or fifth and is scarcely swollen, with the minute sensory
hairs scattered over the inner surface from the base almost to the
apex—not concentrated in a relatively deep and sharply defined
depression.
The above description of the mouth-parts is based upon
dissections of two of the new species (D. fallidihalter and D. flavd)
described herein, and of the European D. obscura , Wtz. The
structure of the various organs is apparently the same in males and
females, and, in our opinion, is such that these midges would be
incapable of piercing the skin. Malloch (1915), however, states that
D. {Pseudoculicoides) griseus , Coq., ‘was taken biting, on the bank
of Sangamon River,’ and that D . cinctus , Coq., has been ‘ recorded
as biting human beings.’ In Accra, specimens of Dasyhelea spp.
occasionally alighted on the arm, but; so far as observed, they made
no attempt to bite; possibly, therefore, the statement regarding
D. griseus may have been due to a similar occurrence, and may
subsequently require modification. At least, until the feeding habits
of the midges of this genus have been carefully studied, the above
statement must be regarded with reserve.
Antennae : segments of the flagellum in the female shorter and
broader them in Culicoides , gradually lengthening towards the apex,
but with the apical ones not distinctly differentiated from the basal;
last four segments (twelve to fifteen) in the male, elongate, the
twelfth to fourteenth inclusive cylindrical, binodose,* the whorls of
• The apical node is usually less well-developed than the basal node, and the hairs are
shorter and more slender; but there is distinct variation in this respect, and in D.fiava y sp. n.
(see p. 196), the apical whorl is obsolete.
hairs arising from nodal rings situated on the basal and apical thirds
of each segment. In both sexes, but more conspicuously in the
males, the segments of the flagellum exhibit a tesselated or
sculptured appearance, due to the presence of small -chitinous plates,
from the upper portion of each of which a hair arises; on the basal
segments (four to eleven) in both sexes this sculpturing appears to
be limited to the proximal portions ( i.e . 9 the area below the whorls)
of each, where the plates are sharply defined and radially arranged;
on the distal segments the plates are arranged as shown in fig. 2.
a
Fig. 2. Terminal segments of antennae of a — D. fvsciformis (<£) and
b—D. inconspicuosa (J). (x 475 circa.)
Thorax without anterior or posterior pits or depressions;
scutellum with a varying number of strong marginal or sub-marginal
bristles and usually a few (one to six) short hairs.
Wings hyaline, unspotted; the venation very similar to that of
Culicoides , but with the first and third veins less distinctly
separated, so that normally only one interspace can be distinguished;
in some species these veins appear to be completely fused.
Legs. Tibiae without apical spurs, but the fore and hind pairs
with oblique or transverse rows of bristles distally; first tarsal
segments of all legs sub-equal, those of the middle legs not
appreciably elongated. Claws small, equal, simple in the female,
divided at the tips in the male; in both sexes the inner margin may
be somewhat prominent, tooth-like, at the base, and agulate near
the middle.
Abdomen. Dorsum covered with minute flattened spines;
segments one to seven each with two small, clear, thinly chitinised,
rounded areas. Spermatheca (fig. 9, b-f) single, heavily or
moderately chitinised, usually sub-spherical, the commencement of
the duct sometimes chitinised for a considerable distance.
External genitalia of the male * Ninth segment : tergite, apical
lobe-like processes well developed, often projecting posteriorly
beyond the tergite and sometimes bearing a median dorsal
triangular chitinised plate; sternite not excavated, sometime^
produced posteriorly in the middle line (c.f. figs. 10, 11). Forceps :
side-pieces similar to those of Culicoides \ claspers simple or divided,
if the latter the branches variously formed, dissimilar. Harpes with
the proximal portions greatly developed, highly chitinised, and in
some species connected medially to form an uninterrupted, or almost
uninterrupted, transverse bar. The distal portions show varying
degrees of development and chitinisation, and in all "the species,
except one, examined by us, the distal portion—bent sharply and
directed posteriorly—of only one harpe is developed, thus causing
asymmetry;, in the exception referred to ( D . similis , sp. n.) the
distal portions of both harpes, appear to be developed, but are fused
in the middle line forming a conspicuous median structure.
Aedoeagus strongly chitinised, usually broad but of variable* form.
• In regard to the nomenclature of the hypopygial structures adopted in this and previous
studies, Part* II and III (1920-1921), on the Ceratopogonine midges of the Gold Coast,
Mr. F. W. Edwards writes us as follows :—* I now think that your harpes are the same structures
which de Meijere has called gonapophyses in the Limnobiidae, and I was probably right in
homologising them with the parameres of Culicidae. There is one difference, at first sight
fundamental, in that in Culicoides and Limnobiidae (c.g., Eriocerd) the structures are dorsal
to the aedoeagus, adjoining the tenth sterAite (which I consider to be represented by your
spicular strip of membrane), whereas in the mosquitoes they are morphologically ventral to
the aedoeagus. However, it would seem b’kely that this is connected with the atrophy of the
anal chitinisations in Culicoides and the Limnobiidae, the parameres moving dorsally and
fusing in the mid-dorsal line in order to replace, in function, the tenth sternite. In other
insects the parameres are said to be normally lateral in position, so that it is easy to understand
that they might pass either dorsally or ventrally. In any case the important idea to lay hold
of is that the parameres form with the basal plate the first (morphologically innermost) ring
of the genital tube, the second ring being the mesosome which is more or less invaginated into
the first. In Culicoides I believe the structure you call the aedoeagus is really the basal plate
only, and that there is no mesosome. Properly speaking the term aedoeagus should include
in this case also the “harpes." The two small chitinous appendages of the ninth tergite,
of course, represent the tenth tergite.’
PUPA. The pupae of Dasykelea much resemble those of
Culicoides , the chief point of difference being in the reduction of the
first and elongation of the second abdominal segments, the shape
and structure of the respiratory trumpets and the arrangement of
tubercles on the body.
The structure of the trumpets, as shown in the species examined
by us, is of especial interest since forms occur which represent
transitional stages between the simple type, as exemplified by
Culicoides , and the complex type as found in at least some species
of Forcipomyia (e.g., F. ingranti , Cart., 1919). Such forms are
shown in fig. 3, b-d \ the last type figured evidently represents
Fig. 3. Pupal trumpets of b — b. JUtviformis ; c — D. tngrofusca ;
d — D.ftuca . {b—c x 360 circa \ d X 220.)
a stage in development immediately preceding that occurring in
Forcipomyia> for by a simple folding back of the distal leaf-like
portion an organ very similar in structure to the trumpet of
F. ingrami could be produced.
i8 3
The chief differences in the arrangement of the body tubercles
is as follows:— Cephalothorax : anterior dorsal tubercles absent,
anterior dorso-median tubercles present (as in Stiiobezzia)t dorsal,
postero-dorsal, and ventro-median tubercles absent but sometimes
represented by minute hairs; the dorsal surface of the thoracic
portion is thus entirely devoid of distinct tubercles, but is, on the
other hand, usually pigmented or striated in circumscribed areas.
Abdomen : tubercles flattened and flange-like, the dorsal antero-
submarginal tubercles small or rudimentary, one only on each side;
the postero-marginal tubercles are situated, as in Stilobezzia , near
the middle transverse line of each segment, around which they form
an almost continuous (interrupted in the 'middle line of the dorsal
and ventral surfaces) band; on segments three to seven they are nine
in number, of which the three central ones (ventro-lateral tubercles)
are the most highly developed. In some species small, rounded,
pigmented areas, arranged in the form of a triangle, are present on
both dorsal and ventral surfaces.
The terminal processes are broad and flattened, and are directed
dorsally and laterally; at the base of each are two prominent
tubercles bearing hairs and spines.
Larva. The mature larva differs from that of Culicoides
chiefly in regard to the size and shape of the head, the position of
the eyes and in the possession of a small hooked pseudopod at the
posterior extremity of the body (fig. 9, g ).
The head is distinctly larger (almost twice as large as in
Culicoides) and tapers more strongly towards the apex, being
obovate rather than sub-conical; it bears minute hairs, only. The
eyes are reniform and are situated far forwards, near the anterior
third. The mouth-parts are similar in structure to those of
Culicoides , but, in the three West African species observed, appear
to be rather more highly developed in so far as the hypopharynx is
larger and more complex, the mandibles are tridentate and the
labium is more highly chitinised and moxe strongly serrated. The
body is elongate and scantily clothed with very minute hairs; the
anal pseudopod is retractile, small and membranous, and bears two
dorsal and two ventral groups of small but strongly chitinised,
recurved hooks which are directed anteriorly. The anal papillae
are short, bifid, with slightly blunted extremities.
184
Specific Descriptions.
Dasyhelea pallidihalter , sp. n.
Measurements.
Female.
Male.
Length of body* .
...
. 1-3 mm.
1*3 mm.
Length of wing ..
...
..o*8 mm.
o-8 mm.
Greatest breadth of wing ...
...
... ... o*3 mm.
0*3 mm.
Head dark brown, clothed with dark hairs. Eyes narrowly
separate in both sexes. Proboscis and palpi brown, the latter with
the third segment uniformly and slightly swollen and almost twice
as long as the fifth; the second, fourth and fifth short, sub-equal.
Antennae dark brown with dark brown hairs; in the female the
segments of the flagellum are sub-spherical at the base, gradually
lengthening, becoming oval, towards the apex, the fourth to eleventh
each bearing a pair of large, strong, bluntly rounded spines; in the
male, segments four to eleven are sub-spherical to ovoid in shape,
the length from two-thirds to one and one-third times the greatest
breadth, segments twelve to fifteen are elongate, about three times as
long as wide, the last broader and pointed distally, the twelfth to
fourteenth sub-cylindrical, each distinctly binodose. Thorax dark
brown with small yellowish-brown areas in the humeral angles and
over the roots of the wings. Scutellum dark brown, becoming
brownish-yellow centrally, bearing four centro-marginal and two
lateral bristles, and a median transverse row of about six short hairs.
Post-scutellum and pleurae dark brown. Wings in the female well
clothed with decumbent hairs which extend almost to the base
between the fourth and fifth veins and are numerous in the anal
angle and fork of the fifth vein. Costa and first and third
longitudinal veins not extending to the middle of the anterior
border, terminating above or slightly beyond the bifurcation of the
fifth vein and enclosing a small but distinct cell or interspace; in the
male less hairy (no hairs at the base of the wing, and very few, if
any, in the anal angle or fork of the fifth vein), the costa terminating
slightly before the bifurcation of the fifth vein. Halteres white, or
almost white. Legs uniformly dark brown. Claws in both sexes
with the inner margin at the base somewhat produced, and near
the middle with a small, angulate prominence. Abdomen : dorsum
• In ail cases taken from the anterior margin of the thorax to the tip of the abdomen of
specimens mounted in carbolic.
i8 5
dark brown, sparsely clothed with dark brown hairs, venter rather
paler brown. Spermatheca highly chitinised, spherical (diameter
50^1) with a conical posterior extension (length about 20 fi) leading
to the duct.
Hypopygium (fig. 4). 'Ninth segment : tergite short and broad,
relatively strongly chitinised and sparsely clothed with moderately
long, strong hairs, posterior margin straight, produced on each side
into a small, rounded, hairy knob, apical lobe-like processes well
developed, often extending as a centrally indented membrane
considerably beyond the extremity of the tergite; stemite broad,
Fio. 4. D. pallidibalter, sp.n., male hypopygium (ventral view), x 400 circa.
produced centrally and posteriorly into a relatively large, more or
less oval, lobe-like process. Forceps : side-pieces well chitinised and
with long hairs, as in Culicoides ; claspers long and slender, rather
strongly chitinised, the basal half pubescent with a few strong hairs,
the apex pointed, slightly depressed, bearing two or three short,
delicate hairs. Harpes as shown in fig. 4, the proximal portions
very strongly chitinised, the distal portion—only that of the left*
harpe is developed—very feebly chitinised, narrow and bent almost
at a right angle to the proximal portion. Aedoeagus somewhat
V-shaped, relatively short, the lateral arms with outwardly directed,
• In this and all subsequent species in which the harpes are not symmetrically developed,
the tide of the insect from which the distal portion arises is stated. The figures of the
hypopygium, however, show the ventral aspect and therefore in them the right side of the insect
lies to the left of the reader.
pointed, beak-like, distal processes and connected anteriorly by a
slightly curved, centrally thickened chitinous bar.
PUPA. Length vg mm. Integument pale brownish-yellow, not
infuscated in any part or heavily shagreened except on the operculum
and on the posterior abdominal segments. Respiratory trumpets
narrow and strongly curved, almost semi-circular; length about
o’3 mm. The trumpets are obliquely and irregularly ringed, the
tracheal trunk in each gives off a number of short lateral branches,
which lead to small elevations on the surface, and terminates distally
in five or six short, blunt processes. Cephalothorax : dorsum of
thoracic portion not pigmented, smooth except on anterior border,
and with a few puckered or striated areas. Anterior marginal tubercle
large, shagreened, bearing a short bristle; dorso-lateral tubercle
rounded, smooth, bearing two or three small hairs, anterior dorso-
median tubercle small, bearing a short hair; ventro-lateral tubercle
moderately large, rounded, bearing a short spine and a short hair;
ventro-median tubercle represented by two minute hairs. Between
the anterior marginal tubercles are two (one dorsal and one ventral)
rounded, unarmed median humps. Dorsal tubercles absent, but
anteriorly represented by one or two minute hairs. Abdomen :
integument reticulated and slightly shagreened at the sides and base
of each segment, more heavily shagreened on the distal segments,
the last entirely shagreened. Tubercles arranged as mentioned on
page 183, the antero-submarginal tubercle small armed with a
short spine, the dorsal and ventral postero-marginal tubercles
apparently unarmed, low, broad and flange-like, the ventro-lateral
postero-marginal tubercles larger and conical, the middle bearing a
short hair, the others a short spine. Terminal processes broad and
flattened, directed dorsally and laterally, each with two posterior
tubercles, bearing respectively a hair and a spine.
Larva. Length about 3 mm. Head obovate, convex dorsally,
flattened ventrally; length 0*2 mm., greatest breadth 012 mm.
Hairs scanty, minute. Mental plate crescentic, moderately
chitinised, apparently bearing nine teeth on each side, the last large
and broadly rounded, the others small, narrowly rounded and closely
apposed. Mandibles large, highly chitinised, the distal portion
bearing three large teeth on the inner side. Hypopharyngeal
sclerite very highly chitinised posteriorly, the posterior portion
*87
armed with three rows of teeth and with a delicate, serrated, fringe-
like expansion on each side. Body scantily clothed with minute
hairs, the last segment with apparently eight pairs of very small
hairs near the extremity. Posterior pseudopod (fig. 9, g) small, the
dorsal groups each consisting of two hooks, the ventral groups each
of four hooks.
Habitat : Nsawam, Gold Coast. Reared from rotting material
collected from the base of a banana plant; March to May, 1920.
Dasyhelea fusciscutellata , sp. n.
Measurements.
Female.
Male.
Length of body (two specimens) ...
i*o mm.
i-i mm.
Length of wing
o*8 mm.
o*8 mm.
Greatest breadth of wing.
. o*3 mm.
0*3 mm.
Head dark brown with almost black
hairs. Eyes
narrowly
separate in the female, very narrowly separate in the male. Clypeus,
proboscis and palpi brown; third segment of the latter cylindrical,
slightly and uniformly swollen, about as long as the fourth and fifth
segments together. Antennae brown with black hairs and, except
on the last five segments, long spines; in the female the segments
ovoid and sub-equal in size gradually elongating towards the apex,
but the thirteenth and fourteenth segments slightly longer than the
eleventh and twelfth, last segment the longest, not produced distally
into a stylet; in the male, segments four to eleven spheroidal,
gradually becoming narrower and more elongate apically, the last
four segments longer, the twelfth and thirteenth sub-equal in length,
each slightly longer than the fourteenth, the last segment about as
long as the penultimate and without a terminal stylet, segments
twelve to fourteen distinctly binodose. Thorax dark brown with
black hairs, the anterior lateral angles sometimes of a pale brown
colour. Scutellum dark brown with four centro-marginal bristles,
two lateral bristles, and two small hairs. Postscutellum and pleurae
dark brown. Wings sparsely clothed with decumbent hairs which
extend basally between the fourth and fifth veins beyond the anterior
cross vein. In the female the costa terminates near the middle of
the anterior border but extends well beyond the bifurcation of the
fifth vein, and the first and third veins form distally a somewhat
indefinite cell; in the male, the costa ends before the bifurcation of
188
the fifth vein and the first and third longitudinal veins form a
distinct cell. Halteres with tawny-yellow knobs and dark brown
stems. Legs brown, uniformly coloured; claws with a basal hairy
extension, of the female simple, each with a slight sub-central
projection, of the male bifid distally, each with a distinct sub-central
projection. Abdomen dark brown with black hairs. Spermatheca
(fig. 9, f) heavily chitinised, somewhat reniform, the anterior portion
ovoid (length 46/u, breadth 49/1), the posterior portion (length 18/O
narrower and sharply curved.
Hypopygium (fig. 5). Ninth segment : tergite broad, slightly
tapering posteriorly and bearing a few moderately long hairs
dorsally, posterior margin without a central notch and with the
lateral finger-like processes small, apical lobe-like processes well
Fig. 5. D. fusciscutellata, spTn., male hypopygium (ventral view), x 400 circa.
*
developed and with a median triangular dorsal chitinised plate
between them; sternite with a somewhat rectangular median
posterior projection. Forceps : side-pieces strongly chitinised, with
long hairs; claspers well chitinised, the basal halves pubescent and
bearing one or two relatively stout hairs, the distal extremities
slightly depressed. Harpes : basal portions strongly chitinised,
broad laterally, narrowing towards the middle line; from the left
basal portion a feebly chitinised, slightly curved, pointed, blade-like
process projects posteriorly as far as the middle of the side-pieces.
Aedoeagus : median transverse basal portion not very broad, heavily
chitinised and slightly twisted and bevelled laterally, lateral portion
189
strongly chitinised, the distal extremities hook-like and directed
dorsally.
Habitat : Nsawam, Gold Coast. Reared from materials collected
from the bases of banana plants, March, 1920.
Dasyhelea similis , sp. n.
Measurements. Female. Male.
Length of body .I*i mm. i*o mm.
Length of wing .07 mm. 07 mm.
Greatest breadth of wing.. ... 07 mm. o*2 mm.
Head dark brown with dark brown hairs. Eyes narrowly
separate in both sexes. Clypeus, proboscis and palpi brown, the
third palpal segment slightly but distinctly swollen at the base and
attenuated distally, in the female about as long as the fourth and
fifth segments together, in the male relatively shorter, about as long
as the fifth. Antennae brown with brownish-black hairs and, except
on the last five segments, long, clear spines; in the female the
segments, with the exception of the last, which is the longest but
carries no stylet, are all sub-equal in size and ovoid in shape, any one
of the apical segments (twelve to fourteen) being, if anything,
slightly shorter than any one of the five (seven to eleven) preceding;
in the male, segments four to eleven spheroidal gradually becoming
narrower and longer towards the apex, the last four segments
elongate, twelfth and thirteenth sub-equal, longer than the others,
twelfth to fourteenth distinctly binodose. Thorax dark brown, the
anterior lateral angles sometimes paler brown. Scutellum dark
brown with four* centro-marginal and two lateral bristles, and two
or more small hairs. Post-scutellum and pleurae dark brown.
Wings sparsely clothed with decumbent hairs which scarcely extend
basally beyond the cross-vein; in the male the costa extends slightly
beyond the middle of the wing, its end lying above the fork of the
fifth vein, and the first and third veins form a distinct cell; in the
female the costa extends beyond the middle of the wing and
terminates distally to the fork of the fifth vein, forming with the
first and third vein a somewhat indefinite cell. Halteres with pale
yellow knobs and dark brown stems. Legs uniformly brown.
Female claws simple, male claws bifid with a slight indication
of a sub-central projection; in both sexes with a hairy extension
• The two middle centro-marginal bristles may be replaced by a single central bristle in
this and other species of Dasybelea.
190
at the base. Abdomen dark brown clothed with dark hairs,
spermatheca (fig. 9, e) sub-spherical (diameter 46/1), moderately
chitinised, the duct scarcely chitinised at its commencement.
Hypopygium (fig. 6). Ninth segment : tergite scarcely tapering
posteriorly, the margin straight with rudimentary finger-like
processes each bearing two setae, the apical lobe-like processes large;
stemite not produced posteriorly. Forceps : side-pieces clothed
with long hairs, claspers strongly chitinised, the basal halves
pubescent, the apical halves bearing one or two short~hairs near their
distal extremities which are depressed. Harpes: basal portions
curved, ribbon-like, symmetrical, joined medially, forming a
Fm. 6 . D. similis , sp.n., male hypopygium (ventral view). X 400 circa.
transverse bar from the centre of which a heavily chitinised structure
(representing the fused distal portions) extends posteriorly and ends
in a rather blunt point. Aedoeagus : median transverse basal portion
strongly chitinised, narrow, somewhat thickened in the middle
(triangular in some views), and but slightly curved, lateral portion
terminating in moderately sharp points and bearing dorsally a
strongly developed barb which projects transversely and is curved
in an anterior direction.
HABITAT : Nsawam, Gold Coast. Reared from rotting material
collected from the bases of banana plants, March, 1920.
D . similis closely resembles D. fusciscutellata but is of a rather
lighter colour, generally with lighter yellow halteres; the species
may be readily distinguished by the form of the spermatheca in the
females and the structure of the hypopygium in the males.
l 9 l .
Dasyhelea luteoscutellata , sp. n.
Measurements.
Female.
Length of body (one specimen) .
. I-l mm.
Length of wing .
.o*8 mm.
Greatest breadth of wing.
.o*3 mm.
Head dark brown with dark hairs.
Eyes narrowly separate.
Clypeus, proboscis and palpi brown, the third palpal segment
longest, slightly swollen basally, the fourth and fifth segments
together longer than the third. Antennae brown with dark brown
hairs and long spines, curved and somewhat hair-like, on segments
four to ten, inclusive; the latter segments sub-spherical to ovoid,
elongating towards the apex, the terminal five segments distinctly
more elongate, together longer than segments three to ten united,
the last segment without a stylet. Thorax dark brown with dark
hairs. Scutellum bright yellow with four centro-marginal and two
lateral bristles, and a small central hair. Post-scutellum and pleurae
dark brown. Wings similar to those of D. pallidihalter . Halteres
with white knobs and dark brown stems. Legs brown, uniformly
coloured. Claws simple without a sub-central prominence.
Abdomen dark brown clothed with dark brown hairs. Spermatheca
sub-spherical (diameter 43/t*); the duct chitinised for a short distance
only.
HABITAT : Nsawam, Gold Coast. Reared from material collected
from the base of a banana plant; the pupa and larva were not
identified.
This species is a small, dark brown or almost black midge,
resembling D. pallidihalter but with the scutellum entirely bright
yellow, not darkened laterally; the long spines on the basal
segments of the antennae are more slender and hair-like than in
D, pallidihalter .
Dasyhelea inconspicuosa , sp. n.
Measurements.
Female.
Male.
Length of body (two specimens) ...
. i*o mm.
i*l mm.
Length of wing .
. 0-7 mm.
o*9 mm.
Greatest breadth of wing.
. o*4 mm.
o*5 mm.
Head dark brown. Eyes broadly contiguous in
the female,
arrowly separate in the male.
Proboscis and palpi
brown, the
192
latter relatively long, the third segment elongate, cylindrical, as long
as the fourth and fifth segments together, the second segment about
half the length of the third. Antennae brown with dark brown hairs;
in the female, segments four and five sub-spherical, six to fifteen
oval, gradually becoming longer towards the apex, the last segment
broad without a stylet (fig. 2, b ); in the male, segments four to eleven
spheroidal to ovoid, segments twelve to fifteen elongate, the twelfth
and thirteenth sub-equal, longer than the fourteenth and fifteenth,
the last segment rather shorter than the fourteenth, without a stylet.
Thorax dark brown with dark hairs. Scutellum yellowish brown
with four centro-marginal and two lateral bristles in both sexes,
small hairs absent. Post-scutellum and pleurae dark brown.
Wings densely clothed with decumbent hairs, which in the female
extend to the base between the fourth and fifth veins. In the female
the costa extends to about the middle of the anterior margin, its
extremity distal to the bifurcation of the fifth vein, forming with the
first and third veins a narrow, slit-like cell; in the male the costa
scarcely reaches the middle of the anterior margin and terminates
above the bifurcation of the fifth vein. Halteres with pale brown
knobs and dark brown stems. Legs brown. Male claws bifid at
their extremities and with a minute sub-basal hairy extension on
their inner sides; female claws simple, with a sub-basal extension
similar to that of the male, but smaller. Abdomen : dorsum dark
brown with brown hairs, venter paler brown. Spermatheca strongly
chitinised, spherical (diameter 32/*), the chitinised portion of the
duct straight (length 5/1).
Hypopygium (fig. 7). 'Ninth segment : tergite scarcely tapering
towards its posterior extremity and scantily clothed with long, stout
hairs, the finger-like processes moderately small and somewhat
pointed, each bearing a relatively long hair near the apex and four
near the base, apical lobe-like processes well developed and bearing
between them a small triangular chitinised plate (not shown in the
figure), stemite prolonged centrally and posteriorly into a long
narrow process which covers the aedoeagus and ends in what looks
like, but actually is not a strongly chitinised spine. Forceps :
side-pieces well chitinised, with very long hairs, claspers with the
basal halves pubescent and the tips depressed, and bearing a few
minute hairs. Harpes\ basal portions heavily chitinised, broad
193
laterally and somewhat L-shaped; distal portion of the left harpe
developed but small, slender and feebly chitinised. Aedoeagus j
median transverse proximal portion slightly curved and tapering
laterally; lateral portions look as if they were twisted, apparently
each composed of two plates, between them a tapering median
gutter-like process with a blunt end (not shown in the figure); the
junction between the plates on each side is very thinly chitinised.
PUPA : Length (two specimens) 2 mm.; dorsum of the cephalo-
thorax somewhat infuscated anteriorly, the anterior margin and the
Fig. 7. />. inconspkuosO) «p.n., male hypopygium (ventral view), X 400 circa.
posterior end of the abdomen conspicuously shagreened. Respira¬
tory trumpets : length about o*2 mm.; curved, and conspicuously
ringed except at the extremities. The main tracheal trunk gives
off laterally, at fairly regular intervals, a few processes which
open on elevations slightly larger than the ordinary corrugations of
the trumpet, and apically about eight short blunt processes arranged
in a fan-like manner. Cephalothorax without strongly armed
tubercles. Anterior marginal tubercles divergent, dark coloured,
and covered with coarse dark spicules bearing a minute terminal
hair; between the anterior marginal tubercles are two small dark
elevations, one behind the other, covered with coaise dark spicules;
anterior dorso-median tubercles triple, each with small setae
internally; dorso-lateral tubercles each showing two slightly rounded
elevations bearing small delicate hairs. Dorsal tubercles represented
by a few minute hairs, and on each side of the middle line are three
dark puckered macules, forming an irregular row across the dorsum;
in front of this row on each side are three similar, if less distinct,
macules. Abdomen : the arrangement of tubercles and form of the
terminal processes are similar to those of the preceding species.
Larva. Length 2*6 mm., greatest breadth o*2 mm. (two
specimens, apparently not quite mature). Head : length about
o*2 mm., greatest breadth 0*14 mm.; well chitinised, conical.
Mental plate resembling that of D . pallidihalter , but apparently
with five rather large coarse teeth on each side in$tead of eight.
Habitat: Oblogo; reared from pupae found in collections of
water in the bottoms of canoes tied to the bank of the river Densu,
also from rotten wood taken from the sides and ends of the same
canoes; December, 1919, to March, 1920. Also larvae, pupae and
adults in the collection of the Liverpool School of Tropical Medicine
obtained from the rotting vegetation at the sides of a water hole at
Christiansborg, Accra, March, 1918.
Dasyhelea nigricans , sp. n.
Measurements.
Length of body .
Length of wing .
Greatest breadth of wing.
Male.
1*3 mm.
1-0 mm.
0*3 mm.
Head very dark brown, almost black. Eyes narrowly separated.
Clypeus and proboscis dark brown. Palpi brown with dark hairs,
third segment the longest, but not so long as the fourth and fifth
together and not appreciably swollen, the last segment distinctly
longer than the fourth. Antennae very dark brown with a
plume of black hairs. Segments four to eleven spheroidal, ^ the
last four segments more elongate, sub-equal in length, without
long spines, the last slightly shorter than the penultimate and
without a stylet; segments twelve to fourteen distinctly binodose.
*95
Thorax dark brown, almost black, with small yellowish-brown
patches at the anterior lateral angles and over the bases of the
wings. Scutellum orange brown in the middle, dark brown at the
sides, bearing four centro-marginal and two lateral bristles, and one
or two median short hairs. Post-scutellum and pleurae dark brown.
Wings sparsely clothed with decumbent hairs. The costa terminates
about the middle of the wing and proximal to the bifurcation of
the fifth vein; the first and third veins form distally a single small
cell. Halteres with yellowish-brown knobs and dark brown stems.
Legs brown, almost uniformly coloured, the femora and two terminal
tarsal segments slightly darker. Claws bifid distally without a
sub-basal projection. Abdomen : dorsum very dark brown, scantily
clothed with black hairs, venter distinctly lighter in colour.
Hypopycium (fig. 8 ). Ninth segment : tergite long, reaching
beyond the ends of the side-pieces, tapering posteriorly and showing
Fio. 8. D. nigricans , ip.n., male hypopygium (ventral view), x 40x5 circa.
a slight concavity at its posterior margin, clothed dorsally with a
few moderately long hairs, finger-like processes well developed,
dark, each with a rounded apex bearing a small terminal hair;
stemite prolonged posteriorly into a cone-like process. For ceps :
side-pieces prolonged distally on their inner sides into sub-conical,
thinly chitinised lobes which are clothed with short, stout hairs—the
196
inner margin from the base of each side-piece as far as the origin
of the lobes is very heavily chitinised; claspers double, the inner
piece more heavily chitinised and shorter than the outer, pubescent,
ending in a strongly pointed process which carries a relatively
stout hair, and showing a secondary barb on its convex margin at
about the apical third, the outer portion less heavily chitinised,
pubescent on the basal half or two-thirds and of normal form, with
the apex depressed and bearing a few small hairs. Harpes : basal
portions rather broad, strongly chitinised, asymmetrical; from the
right basal portion there arises a curved, heavily chitinised, blade¬
like structure which tapers to a point distally, and is prolonged
posteriorly as far as the distal end of the main part of the side-piece;
laterally the basal portions are produced into strongly chitinised
bars which run diagonally backwards and join the end of the
chitinised inner margin of the side-pieces. Aedoeagus : strongly
chitinised, broad; the median transverse proximal portion with
wing-like prolongations, the lateral portions each composed
of a dorsal and ventral plate, the extremities divergent, pointed,
resembling a pair of shears.
Habitat : Accra, Gold Coast. Two males taken in the evening
upon the windows of the laboratory, January and February, 1920.
Dasyhelea flava , sp. n.
Measurements.
Female.
Male.
Length of body (three specimens)
... 1*2 mm.
1*3 mm.
Length of wing
... o*8 mm.
o*9 mm.
Greatest breadth of wing.
... 0*2 mm.
0*3 mm.
Head yellow, the middle of the occiput brown. Eyes separated,
narrowly in the female, more widely in the male. Clypeus
yellow. Proboscis and palpi brown, the first segment of the
palpi very small, the third segment inflated, about one-third
longer than the fifth in the female; third segment of palpi
in male cylindrical, less inflated than in the female. Antennae
brown with dark hairs; in the female, segments four to ten
bear stout sensory spines and are sub-spherical to oval in shape,
the length varying from one to one and one-fifth the width,
segments eleven to fourteen slightly longer, the last segment tapering
to a narrowly rounded apex and almost one and three-quarter times
x 97
the length of the preceding segment; in the male, segments four to
eleven oval to sub-oval in shape (almost hexagonal in transverse
section), varying in length from two-thirds to one and a half times
the width, last four segments gradually decreasing in length, twelve
to fourteen, not binodose but exhibiting the' usual sculpturing,
with a basal whorl of hairs and shorter hairs distally, the last
segment longest, longer than the twelfth, basally very broad
and tapering to a pointed apex. Thorax canary yellow with three
broad longitudinal brown bands: a median, which runs from the
anterior margin to the posterior third of the dorsum and is frequently
divided down its middle by a narrow yellow line, and lateral bands
extending from the anterior fourth of the dorsum almost to the
scutellum. In the male the thoracic pattern is less conspicuous, the
yellow markings being contracted so that the general colour appears
brown with yellow markings, rather than yellow with brown
markings. Scutellum canary-yellow with four centro-marginal and
two lateral black bristles, in both sexes; small hairs absent. Post-
scutellum dark brown. Pleurae almost entirely canary yellow with
small brown markings. Wings hyaline without spots, clothed with
decumbent hairs which extend basally well beyond the cross-vein.
Halteres canary-yellow in both sexes. Legs brown, rather lighter
coloured in the male, the knees infuscated. Claws simple, bifid at the
tips in the male. Abdomen : dorsum dark brown, venter somewhat
tawny, especially towards the apices of the segments. Arising from
the centre of the anterior margin of the eighth stemite in the female
and extending forwards into the seventh segment is a blunt, chitinous
and apparently tubular process (fig. 9, a ); the lower laminae appear
to arise from the anterior edge of the eighth stemite, but the upper
laminae extend backwards internally almost to the edge of the
segment. This structure is conspicuous in specimens which have
been treated with caustic potash, but is largely obscured by pigment
in carbolic preparations; even in dry specimens, however, a small
portion of the lower plate can usually be detected near the
middle of the anterior margin of the eighth segment. Spermatheca
(fig. 9, b ), single, moderately chitinised, sub-spherical to oval
(44A1 by 40 ft) in shape with the beginning of the duct chitinised.
Hypopygium (fig. 10). Ninth segment : tergite broad basally,
tapering distally, sparsely clothed with strong hairs, posterior margin
198
not notched and without finger-like lateral processes, stemite
relatively narrow, with a conspicuous but not heavily chitinised
median pincer-like process projecting backwards ventral to the
aedoeagus. Forceps : side-pieces short and stout, broader apically
than basally, rather scantily clothed with moderately long hairs ;
claspers bifid, highly chitinised, antler-like, the inner branch pointed
and bearing two setae a little beyond the middle, the outer branch
strongly curved, pointed, with a prominent seta near the base.
Harpes : basal portions articulating with the base of the side-pieces,
strongly chitinised; from the right basal portion arises a lightly
chitinised sickle-like process which is very long and tapers
gradually towards the tip—it is first directed dorsally, but later
curves strongly in a ventral direction and distally is covered
with delicate spicules. Aedoeagus : a broadly-ending gutter-like
structure tapering but slightly from base to apex, the median
transverse portion and the lateral portions heavily chitinised.
PUPA. Length (two specimens) vy mm. Respiratory trumpets
long, narrow and curved, arising from small tubercles; length about
199
0*2 mm. Distal extremity not infuscated, slightly broadened and
without rings, middle portion irregularly ringed, base slightly
constricted; lateral openings of the tracheal trunk occur from base
to apex, about six being situated near the' tip. Cephalothorax :
anterior marginal tubercles large, conical, heavily shagreened,
bearing a very short bristle. In the middle line between
the anterior marginal tubercles is an antero-posterior row of three
Fio. 10. D.flaoa, ip.n., male hypopygium (ventral view), x 400 circa.
unarmed somewhat heavily shagreened tubercles, the most
anteriorly placed being slightly ventral, the middle one slightly
dorsal, to the marginal tubercles. Anterior dorso-median tubercle
represented by two or three small hairs; dorso-lateral tubercle some¬
what prominent, irregularly shaped and bluntly rounded, bearing a
moderately long and a short hair; ventro-lateral tubercle bearing a
minute hair; ventro-median tubercle represented by a small hair.
Tfie ventral surface also bears small, highly chitinised, unarmed
tubercles, one on each side of the middle line near the centre of the
cephalic region. Abdomen more heavily shagreened than, but
otherwise similar to, that of D. pallidthalter.
LARVA. Adults of this species were reared from larvae obtained
from banana fibre which had been isolated in small tubes; the larval
200
pelts were not subsequently found, but as no other species of
Dasyhelea was reared from this particular sample of banana fibre
in the course of some weeks’ observation, the Dasyhelea larvae found
therein have been ascribed to D. flava .
Length (three specimens) 2*9 mm.; greatest breadth 0*2 mm.
Head : length 018 mm., greatest breadth 0*12 mm. These larvae
are almost indistinguishable from those of D . pallidihaltef , the only
apparent difference being in the size and form of the teeth on the
mental plate, in D. flava larva the most lateral tooth on each side
is smaller, while the others are distinctly larger and coarser than in
larvae of D. pallidihalter.
Habitat : Nsawam and Dodowah, Gold Coast. Reared from
rotting vegetable material collected at the bases and roots of banana
plants, January to May, 1920.
Dasyhelea juscipleuris, sp. n.
Measurements.
Length of body (one female) . . 1*2 mm.
Length of wing .i*o mm.
Greatest breadth of wing.0*4 mm.
Head brown. Eyes narrowly separate. Proboscis and palpi
brown, the fourth and fifth palpal segments short, sub-equal, third
segment slightly inflated and rather longer than the fourth or fifth.
Antennae brown; segments four to ten inclusive sul>spherical to
broadly oval in shape, in length from four-fifths to one and one-
quarter times the greatest width, segments eleven to fourteen oval
and in length about one and a quarter times the greatest width, the
fifteenth slightly longer and about twice as long as wide. Thorax
yellow, becoming yellowish-brown posteriorly in front of the
scutellum, with broad brown or blackish-brown median and lateral
bands; the median band, which has a shining appearance in certain
lights, is of almost uniform width and extends from the anterior
margin to about the middle of the thorax, the lateral bands are
broad and rounded anteriorly, gradually narrowing posteriorly, and
extend from the anterior fourth of the thorax almost to the scutellum;
each lateral band near the middle of its outer border gives off a
narrow dark stripe which is directed forwards and ends a short
distance behind the antero-lateral angle. Scutellum entirely pale
201
yellowish-brown with three or four centro-marginal and two lateral
bristles; short hairs absent. Post-scutellum dark brown. Pleurae
almost entirely yellowish-brown with a few small darker brown
markings. Wings hyaline, sparsely clothed with decumbent hairs
which scarcely extend basally beyond the cross-vein and in the
anterior apical portion are almost confined to the margin and to the
fold above the upper branch of the fourth vein. The costa extends
slightly beyond the middle of the wing and distinctly beyond the
bifurcation of the fifth vein; the single cell between the first and third
veins distinct, rather longer than broad. Haltcres yellow-brown.
Legs : distinctly light brown with the extreme bases of all the tibiae
and the apices of the Hind tibiae very dark brown; the fifth tarsal
segments of all the legs are somewhat infuscated. Claws simple.
Abdomen : dorsum dark brown, with the hind margins of the
segments narrowly but distinctly yellowish; venter of a rather pale
brown colour. Spermatheca (fig. 9, c) single, oval to pyriform (average
about 61 fi by 42 /*), highly chitinised, the commencement of the duct
chitinised for a short distance (about 8/*)•
HABITAT: Accra, Gold Coast. Two females taken on the arm,
one in a bungalow near a light at 9 p.m., April 18th, 1920, and the
other in the laboratory in May, 1920. Numerous specimens ($ $)
bred later.
A yellowish-brown midge somewhat resembling D. flava in
appearance, though less brightly coloured.
Dasyhelea flaviformis , sp. n.
Measurements. Female. Male.
Length of body . i*o mm. i*o mm.
Length of wing .o*6 mm. 07 mm.
Greatest breadth of wing.o*2 mm. o*2 mm.
Head brown with lighter brown borders round the eyes and
occiput. Eyes narrowly separate. Proboscis and palpi brown,
third segment of the palpi not swollen, longer than the fourth and
about as long as the fifth segment. Antennae brown with dark
brown hairs; segments four to ten in the female with curved spines,
sub-spherical, as broad as long, segments eleven to fourteen rather
longer than broad, the terminal segment the longest, being two and
a half times as long as broad and almost twice as long as the
202
preceding segment; the last four segments of the male elongate,
segments twelve to fourteen binodose. Thorax tawny-yellow with
broad, longitudinal, brown bands arranged in much the same pattern
as those of D. flava and of D. fuscipleuris : a broad median band,
divided by a narrow, median* tawny stripe extends almost from the
anterior margin to the posterior third of the dorsum, and lateral
bands, one on either side, extend from the anterior third of the
dorsum almost to the scutellum. Scutellum tawny-brown with
slightly darker brown sides, carrying three central and two lateral
bristles in both sexes; short hairs absent. Post-scutellum dark
brown. Pleurae tawny with two or three dark brown markings.
Wings hyaline; decumbent hairs rather more scanty than in D. -flava ,
extending basally between the fourth and fifth longitudinal veins to
about the point of separation of the first and third longitudinal veins,
the anal cell bare and the anterior apical region rather more sparsely
clad. Halteres with brown stems and cream-coloured knobs. Legs
Fig. ii. D. fiaviformUy sp.n., male hypopygium (ventral view), x 400 circa.
almost uniformly light brown, with dark knee spots. Claws simple
and equal. Abdomen dark brown; in the female there are semi¬
lunar tawny markings on either side of the middle line on the dorsum
of the fifth, sixth and seventh segments, and the apex of the abdomen
is pale; in the male entirely dark, except for small paler lateral spots
on the first and second segments. Spermatheca (fig. 9, d) pyriform
(length 43 m, breadth 35 m), moderately highly chitinised, the
commencement of the duct chitinised.
Hypopygium (fig. 11) resembling that of D. flava . Ninth
segment : tergite broad basally, narrowing slightly towards the
203
apical margin which is not notched, finger-like lateral processes
rudimentary but bearing a single rather long stout hair; stemite
with the median, pincer-like extension well developed. Forceps :
side-pieces relatively longer than in D. flava , claspers more slender
and less curved, and bearing a rudimentary, conical, second branch
on their inner sides basally, the basal portion only pubescent.
Harpes similar to those of D . flava but relatively larger and stouter;
the sickle-like structure arising from the right basal process is very
long and strongly curved, and apically is devoid of spicules.
Aedoeagus differs from that of D. flava in that the distal extremity
gives off two curved chltinous processes, one on either side of the
middle line, resembling a pair of forceps.
PUPA. Length about 1*5 mm.; somewhat infuscated at the
anterior end, integument shagreened. Respiratory trumpets
(fig-3*6): length about 0*15 mm.; breadth about 15/*. They are
infuscated apically, and are long, narrow and curved, arising from
slight papillae; the main tracheal trunk begins to give off lateral
branches just above its base, the first four widely separate but those
near the tip more closely situated, together forming a series of nine
or ten branches. The trumpets are irregularly ringed in the
middle and show no definite tubercles. Cef halo thorax infuscated
anteriorly; operculum dark, highly chitinised and coarsely
shagreened, with two low and rounded, but relatively large,
elevations in the middle line. Anterior marginal tubercle well-
developed, conical, bearing a very small seta; dorso-lateral tubercles
well-developed, bearing several minute hairs; anterior dorso-median
tubercle almost indistinguishable, being represented by a few minute
hairs; ventral tubercles undeveloped, represented by minute hairs.
Dorsum of thoracic region infuscated, without tubercles but with
numerous macules. Abdomen exhibiting no peculiarities, of the
usual pattern but rather poorly chitinised.
HABITAT : Oblogo, Gold Coast. Reared from rotten wood taken
from a canoe in the river Densu, May, 1920. The pelt of a larva
which had been isolated in a tube with a piece of the rotten wood
in which it was found and from which a pupal pelt and an adult of
this species were obtained, was not recovered.
A small brown midge resembling in markings D. flava , but with
the lighter coloured areas a dull tawny-yellow in place of brilliant
canary-yellow.
204
Dasyhelea fusca , sp. n.
Measurements.
Female.
Male.
Length of body .
... i*5 mm.
1*6 mm.
Length of wing .
. i*i mm.
1*3 mm.
Greatest breadth of wing ...
. o*4 mm.
o*3 mm.
Head brown with brown hairs. Eyes broadly contiguous in the
female, narrowly separate in the male. Proboscis pale brown.
Palpi brown, third segment slightly swollen, distinctly shorter than
the fourth and fifth segments together. Antennae in the female
brown, torus much darker than the segments of the flagellum, with
dark hairs and large, curved spines (in addition to short blunt
spines) on all the segments; segments four to eleven sub-spherical to
oval, constricted apically, the last segment elongate, at least one and
a half times as long as the penultimate, with a distinct stylet. In
the male, strongly plumose, the whorl hairs dark brown; segments
three to eleven spheroidal, segments twelve to fourteen elongate,
strongly binodose; last segment stout and cylindrical, with a
conspicuous stylet. Thorax dark brown with paler brown to
yellowish areas at the anterior lateral angles and bases of the wings
(in fresh specimens the dorsum is greyish-pruinose), sparsely clothed
with dark brown hairs. Scutellum pale yellowish-brown with
five or six centro-marginal and two lateral bristles, arid about
six (four to seven) short median hairs. Post-scutellum and pleurae
dark brown. Wings almost unicolourous but slightly infuscated at
the junction of the first and third veins and costa; surface rather
thickly clothed with long hairs, which extend basally beyond the
cross-vein. First and third veins fused, extending for a considerable
distance parallel to the costa and joining the latter well beyond the
middle of the anterior border and the bifurcation of the fifth vein.
Halteres with cream-coloured knobs. Legs light brown, the
knees slightly darker, the apices of the tarsal segments infuscated,
fore and hind femora each with a dark spot above the
apex. Claws simple, bifid distally in the male. Abdomen dark
brown with brown hairs, which are most numerous at the sides and
on the distal segments. Spermatheca more or less pyriform (length
61 /i, breadth 45/0 in shape and feebly chitinised.
Hypopygium (fig. 12). Ninth segment : tergite with moderately
long hairs dor sally, broad, gradually tapering towards the posterior
205
margin, the finger-like processes well developed, dark, each with a
stout hair at the apex and a group of five or six shorter hairs at the
base; stemite relatively broad, without a central posterior projection.
Forceps: side-pieces short and broad, clothed with short, stout
hairs; claspers pubescent, rather longer than the side-pieces, the tips
slightly rounded and bearing a few minute hairs. Harpes : basal
portions somewhat asymmetrical, large, strongly chitinised, curving
forwards towards their junction near the middle line; distal portion
of the right harpe in the form of a stout, strongly chitinised, blade¬
like structure with a pointed, slightly re-curved apex, extending
posteriorly almost as far as the posterior margin of the ninth tergite.
Aedoeagus large, projecting posteriorly to about the level of the
middle of the side-pieces, broad basally, narrowing slightly apically
as shown in fig. 12.
PUPA. Length 2’3 mm.; relatively large and well chitinised,
with the cephalo-thoracic portion slightly infuscated and with
206
characteristic, loofah-like, trumpets. Integument shagreened.
especially over the middle of the anterior part of the cephalothorax
and on the last abdominal segment. Respiratory trumpets (fig. 3, d) :
length about 0*3 mm., broad and flat, arising from small tubercles;
the basal third cylindrical, dark coloured, annulated, the apical two-
thirds expanded and flattened, with very numerous tracheal branches
arranged somewhat like the leaflets of a long pinnate leaf which has
been doubled upon itself so that the fold is at the distal end and
the base and tip at the proximal end—the branches are apparently
double processes arising from the main tracheal trunk. Cephalo¬
thorax'. anterior marginal tubercles prominent, conical, divergent,
covered with coarse dark granulations and ending each in a double-
pointed process bearing a delicate hair; anterior dorso-median
tubercles somewhat irregularly shaped and bearing small and delicate
setae on their inner sides; dorso-lateral tubercles highly granular,
bearing one small and one minute seta. On each side of the
anterior marginal tubercles is a dark, conical, coarsely granulated
nodule, and immediately dorsal to this a smaller and flatter
granulated nodule. Dorsal tubercles apparently represented by
socket-like marks (which may be surrounded by a dark zone and may
show centrally minute points resembling small spines) and by small
dark puckered macules—there are about eighteen of the latter, of
which fourteen (seven on each side of the middle line) are arranged
in an irregular transverse row extending across the dorsum a short
distance posterior to the trumpets, and the others are situated
anteriorly two on each side of the middle line. Large dark areas,
which appear to be slightly raised, also occur over the bases of the
wing sheaths and on that part of the cephalothorax immediately
posterior to them, and other macules may be distinguisned on
the posterior portion of the dorsum. Abdomen : tubercles poorly
developed; on both surfaces of each segment, but more conspicuous
on the dorsum, are three rounded and darkened macules arranged
in the form of an equilateral triangle with its apex posteriorly
directed.
Habitat : Oblogo, near Accra. Pupae and pupal pelts were
found in stagnant water contained in canoes which were tied to the
bank of the river Densu; and pupae in the rotten wood forming the
sides and ends of these canoes. Larvae were not obtained.
207
Dasyhelca nigrofusca , sp. n.
Measurements.
Male.
Length of body ..
... 1-5 mm.
Length of wing ... .
... i*o mm.
Greatest breadth of wing.
... o«3 mm.
Head dark brown, almost black. Eyes broadly contiguous
dorsally. Proboscis and palpi dark brown, the third segment of the
palp distinctly swollen and rather longer than the fourth or fifth,
which are sub-equal in length. Antennae dark brown with almost
black hairs; segments four to eleven more or less hexagonal, in length
from three-fourths to rather more than once the width, segments
twelve to fourteen elongate, binodose, decreasing in length from about
three times to two and a half times the width, terminal segment
broad, rather shorter than the twelfth segment and with a small
apical stylet. Thorax dark greyish-pruinose with sepia markings,
the lateral margins narrowly yellowish; scantily clothed with dark
hairs. Scutellum dark brown in the middle, yellow laterally, with
a row of eight sub-marginal bristles and about half a dozen
median short hairs. Post-scutellum dark brown. Pleurae yellow with
dark brown markings. Wings hyaline, well clothed with decumbent
hairs. First and third veins apparently completely fused, the
bifurcation of the fifth vein lying immediately below the apex of the
costa. Halteres brown with paler coloured but slightly infuscated
knobs. Legs conspicuously banded, femora dark, slightly paler
basally and with a distinct pale band before the apex, knee joints
dark, tibiae with dark bands at the base, middle and apex, tarsal
segments paler, with indications of dark bands at the apices;
clothed with dark hairs. Claws simple and equal, bifid at the tips.
Abdomen sepia-coloured, clothed with dark brown hairs; venter
yellowish.
Hypopygium (fig. 13). Ninth segment : tergite scarcely tapering,
the posterior margin straight, the finger-like processes well
developed, each bearing a single hair and with a prolongation
of the tergite connected with them dorsally; stemite without
a central posterior extension. Forceps : side-pieces of the
usual form; claspers as long as the side-pieces, the basal halves
pubescent and carrying two short hairs near the middle and three
or four short hairs near the apex. Harpes : basal portions strongly
208
chitinised and bent in aft anterior direction, that of the right side
being shorter and more abruptly bent and articulating with a double
curved, blade-like distal process; the latter is very long and reaches
posteriorly as far as the apices of the side-pieces, it is broader in the
middle than at either extremity, and is serrated at the apex.
Aedoeagus broad basally, narrowing somewhat apically, the distal
portion consisting of two highly chitinised hook-like rods on either
side connected by a ventral wall of thin chitin; these lateral rods are
not on the same plane, and the .ventral pair are longer, less sharply
bent medially, and notched at the ends.
Fig. 13. D. nigrofusca , »p.n., male hypopygium (ventral view), x 400 circa.
PUPA. Length 2*6 mm.; shagreened but not highly chitinised.
Respiratory trumpets (fig. 3, c ): length about 0*1 mm., short,
broad and constricted at the base, arising from small tubercles
and with the basal half covered with minute squamose spines; lateral
processes (numbering about twenty) of the tracheal trunk given off
at regular intervals—at first in a single, later in a double row,
extending from a short distance above the base to the apex.
Cephalothorax : spines and tubercles poorly developed; operculum
209
wanting. Anterior marginal tubercle small, bearing a small bristle;
dorso-lateral tubercle bearing two small hairs; anterior dorso-median
tubercle very small with two hairs; ventro-median tubercle minute,
bearing a small hair; ventro-lateral tubercle represented by two or
three small hairs. Dorsum of the thoracic portion without tubercles
or spines. Abdomen : integument with transverse cellular
sculpturing and minute spicules basally and laterally; these spicules
are most numerous and most highly developed on the last segment
and its terminal processes. The segments are furnished with the
usual backwardly projecting shelf-like tubercles, those forming the
ventro-lateral series being the most prominent.
HABITAT: Dodowah, Gold Coast (north-east of Accra), reared
from materials collected from a rot-hole in a mango tree, March 7th,
1920. The larva was not identified.
Dasyhelea fusciformis , sp. n.
Measurements.
Length of body .
Length of wing .
Greatest breadth of wing.
Female.
1*3 mm.
i-i mm.
0*3 mm.
Head ,brown with brown* hairs. Eyes very narrowly separate,
almost contiguous. Clypeus, proboscis and palpi brown; all
segments of the latter somewhat swollen, the third cylindrical, longer
than the fourth and fifth segments together, the fifth much shorter
than the fourth. Antennae brown with brown hairs and long curved
spines on all the segments; segments of the flagellum sub-spherical
to oval in shape, each with an apical constriction, the last segment
elongate with a distinct stylet (fig. 2, a ). Thorax dark brown with
dark hairs, the anterior-lateral angles and areas at the bases of the
wings paler brown. Scutellum light brown with eight centro-marginal
and two lateral bristles, and three short hairs. Post-scutellum and
pleurae dark brown. Wings: decumbent hairs extending to the
base of the wing between the fourth and fifth veins; costa reaching
the middle of the anterior border and terminating beyond the
bifurcation of the fifth vein, the first and third veins distally forming
a small cell. Halteres with cream-coloured knobs and dark brown
stems. Legs uniformly brown, the claws simple. Abdomen dark
brown, lighter coloured distally, with dark brown hairs.
210
Spermatheca sub-spherical (length 53/1, width 48/i), moderately
chitinised, the duct wide at its origin, tapering gradually and
chitinised for a very considerable portion (50/1) of its length.
PUPA. Length 2 mm., moderately well chitinised, the integu¬
ment shagreened and the last abdominal segment with small
spicules. Respiratory trumpets : length about 0*17 mm., breadth
0*03 mm., rather short, broad and almost straight, with a slight
basal constriction and with the surface covered with minute,
imbricated, flattened spines. The tracheal trunk gives off three or
four lateral branches at regular intervals in the basal three-fourths
of the trumpet, and a series of about a dozen close together in the
apical fourth. Cephalothorax : anterior marginal tubercles wanting
—the operculum missing from the single pelt examined; anterior
dorso-median tubercle poorly developed, bearing a single small hair;
dorso-lateral tubercle small, carrying two short hairs; ventro-lateral
tubercle indistinguishable; ventro-median tubercle represented by
two small hairs. Integument of the dorsum of the thoracic portion
with puckered macules, but without tubercles. Abdomen with
tubercles of the usual type and arrangement.
The pupal stage is apparently short, as the adult insect appeared
on the fourth day after isolating the larva. The pupal pelt was
found with the cephalothorax protruding from a piece of rotten
wood.
LARVA. A single larva of this species was collected and isolated
in a small tube; it was rather large, with well-formed hooks at the
posterior end of the abdomen. Its movements were slow and
deliberate and it was observed to crawl up the sides of the glass
vessel in which it was contained, but afterwards it quickly buried
itself in the rotten wood and was not seen again. After the insect
had hatched the pupal pelt was secured, but the only portion of the
larval pelt found was the head; this was dark-coloured and heavily
chitinised, and measured 0*3 mm. in length and 0 2 mm. in greatest
width.
HABITAT: Oblogo, Gold Coast. Reared from materials collected
from a rot-hole in a tree ( Cynometra sp., probably C . megalophylla ,
Harms.), May, 1920.
D . fusciformis is a brown midge closely resembling D . fusca, but
smaller in size and with the colouring of the scutellum and the lighter
patches on the thorax pale brown rather than yellow. The
spermathecae and pupae of these two species are quite distinct.
The species of Dasyhelea described above may be separated as
follows: —
Females
1. Antennal segments sub-spherical or oval, not constricted apically,
the last segment not produced into-a stylet; scutellum with at
most four centro-marginal bristles ... ... ... ... 2
Antennal segments constricted apically,' the last with a distinct
stylet ; scutellum with at least five centro-marginal bristles ... 9
2. Species mainly dark brown in colour.. . 3
Species yellowish with dark brown markings. ... 7
3. Scutellum dark brown, at most with the central portion yellowish... 4
Scutellum entirely yellowish ... ... ... ... ... ... 6
4. Scutellum yellowish in the middle ; halteres white, pallidih alter, sp. n. (p. 184)
Scutellum entirely dark brown; halteres yellowish ... . 5
5. Spermatheca reniform ;* claws with a distinct sub-central pro¬
jection* . fusciscutellata , sp. n. (p. 187)
Spermatheca spherical; daws without a distinct sub-central pro¬
jection ... ... similis , sp. n. (p. 189)
6. Scutellum bright yellow ; halteres white ... lutfoscutellata, sp. n. (p. 191)
Scutellum yellowish-brown ; halteres yellowish-brown .
. inconspicuosa , sp. n. (p. 191)
7. Species bright yellow ; the eighth abdominal segment with a ventral
backwardly directed chitinous process . Hava, sp. n. (p. 196)
Species dull yellow ; the eighth abdominal segment without such a
process ... ... ... ...,, ... ... ... ... 8
8. Larger species (wing length i*o mm.); halteres yellowish-brown ;
hind margins of abdominal segments narrowly yellowish
... .*.. ... fuscipleuris , sp. n. (p. 200)
Smaljer species (wing length 07 mm.) ; halteres cream-coloured ;
semilunar tawny markings on dorsum of fifth, sixth, and seventh
abdominal segments . Haviformis^ sp. n. (p. 201)
9. Spermatheca pyriform, feebly chitinised 5 scutellum with five or six
centro-marginal bristles ... ... ... ... fusca, sp. n. (p. 204)
Spermatheca spherical, moderately chitinised, with, a very long
posterior extension ; scutellum with eight centro-marginal
bristles . fusciformis, sp. n. (p. 209)
Males (separation on hypopygial characters)
1. Claspers single .
Claspers double.
2. Ninth sternite with a central posterior extension
Ninth stemite without a central posterior extension
2
6
3
4
3. Posterior extension of ninth sternite large, ob-ovate .
. paJltdibalUr , sp. n. (p. 184)
Posterior extension of ninth sternite smaller, somewhat rectangular
. *. jusciscuuUata, sp. n. (p. 187)
Posterior extension of ninth sternite cone-like inconspicuosa , sp. n. (p. 191)
4. Distal portions of harpes fused, forming a single median process ...
... ... ... ... ... ... ... stiHiltS) sp. n. (p• 189)
Distal portion of one harpe only developed. 5
5. Aedoeagus relatively short, reaching about one-third the length of
the side pieces ; distal fourth of the harpal process attenuated,
the apex serrated ... . nigrofusca , sp. n. (p. 207)
Aedoeagus long, reaching beyond the middle of the side pieces;
harpal process not attenuated distally, the apex pointed
•.. ... ... ... ... ... ... fusca, sp. n. (p. 204)
6. Inner portion of clasper small, conical ... flavijormis, sp. n. (p. 201)
Both portions of clasper large and well developed. 7
7. Central posterior extension of the ninth sternite cone-hke ...
. nigricans , sp. n. (p. 194)
Central posterior extension of the ninth sternite pincer-like
. Hava, sp. n. (p. 196)
REFERENCES
Carter, H. F. (19x9). New Wcit African Ceratopogoninae. Ann. Trop. Med. & Parasite
Vol. XII, pp. 2S9-302.
Caster, H. F., Ingram, A., and Macro, J. W. S. (1920). Observations on the Ceratopogonine
Midges of the Gold Coast with descriptions of new species. Part II: Ibid., VoL, XIV,
pp. 211-274. Part III: Ibid., pp. 309-331.
Kievter, J. J. (1917). Chironomides d’Am£rique conserves au Mus6e National Hongrois de
Budapest. Ann. Mm. Nat. Hung, Vol. XV, pp. 292-364.
-(1919). Chironomides d*Europe conserves au Mus6e National Hongrois de Budapest.
Ibid., Vol. XVII, pp. 1-160.
- (1919). Observations sur les Chironomides (Dipt.) d£crit» par J. R. Malloch. Bull.
Soc . Ent. de France, No. ro (May), pp. 191-194.
Malloch, J. R. (19x5). Some additional records of Chironomidae from IHinois and notes
on other Illinois Diptera. Bull. Ill . State Lab. Nat. Hist., Vol. XI, pp. 305-363.
213
THE PREVALENCE AND CHARACTER
OF TUBERCULOSIS IN HONGKONG
BY
HENRY HAROLD SCOTT
GOVERNMENT BACTERIOLOGIST, HONGKONG
(Received for publication I July , 1921)
I. GENERAL CONSIDERATIONS
The statement has been made that not only is tuberculosis a
common disease in the tropics, but that it appears to be increasing
in its ravages in the East. Seeing that the treatment of tuber¬
culosis, both prophylactic and curative, consists largely in fresh air
and sunshine, it seems strange that this disease should be rife in
tropical countries where sunshine is so prevalent and living in the
open air is the rule.
A very brief experience as medical officer in charge of the
mortuary sufficed to show that tuberculosis was a frequent cause of
death in Hongkong, and I deemed it an investigation well worth
undertaking to determine the varieties of the disease as met with
here, the portals of entry, the mode of spread, and, if possible, to
determine the primal cause or causes responsible for the condition
and those aiding its dissemination, with a view to elucidating
measures for its prevention.
I have excluded those cases which showed merely signs of
old-standing tuberculosis, for previous investigations have proved
that after middle life nearly all bodies yield evidence of some healed
focus of the disease. A particularly noticeable feature in my series is
the large percentage of children of tender age who die from tubercu¬
losis. Another point worthy of note is the fact that whereas at home
intestinal tuberculosis as a primary condition is fairly common in
children and is ascribed to the drinking of infected milk, in tropical
countries primary intestinal tuberculosis is met with although milk
214
is not taken as a regular food; this holds good here also, though
it is only right to add that primary intestinal ihfectibn appears to
be comparatively rare in Hongkong.
Before proceeding to speak in detail of the cases dealt with in
this investigation, it will not be out of place to make a few general
remarks on the disease. As is well known, the bacillus may persist
for a long time in dust, in dried sputum, in urine, and so forth.
When one sees how large a number of patients exhibit signs of
tuberculosis after death, several of whom showed no marked signs
during life, and when, as we know, the bacilli are shed broadcast
not only in the sputum but in the fine spray of coughing and
sneezing consumptives, the ppportunities for the organism to gain
entrance and to propagate are many* .Thus, the bacillus has been
found in the sweepings of rooms (Manfredi at Naples, Marpmann
at Leipzig), on fruit exposed for sale (Schnirer), in the wards of
hospitals (Strauss, Le Noir), in books, in the bodies of flies
(Nuttall, Andr6), on clothes (Josephson, Notel, Chausse), as quoted
by Etienne Burnet: ‘ Ou trouve comme une quintessence des
poussieres atmospheriques dans ces poussi&res fines que Ton pompe
par le vide (vacuum cleaner); avec elles s’est depose ce qu'il y a de
plus 16 ger dans ce que soulfevent nos semelles et Tintense circula¬
tion de nos rues, et ce que crache une population malheureusement
trop insouciante.*
As stated by Del6pine, the difficulties of elucidating the causes
of an infectious disease depend on the fact that we must take account
of the various ways in which the predisposing and determining
factors have combined in their action, and the dijficulties are
increased when we attempt to assign to each its appropriate share.
He groups these factors under the following heads: (i) Distribution
and habits of the pathogenic organism. (2) Conditions influencing
its number. (3) The pathogenicity and toxicity of it, or, in other
words, the conditions influencing its virulence. (4) The oppor¬
tunities which the organism may have of gaining access to one or
more channels of infection. (5) The conditions influencing the
resistance which any possible host may be capable of opposing to
the attacks of the organism. (6) The frequency and degree of
completeness of recovery from occult or manifest infection. This
comes into play more with regard to the question of adult tuber-
215
culosis, whereas the present investigation is concerned with the
disease as found in children.
Of these factors those of special importance are the number and
virulence of the organism on the one side and the degree of
resistance which the subject attacked may be able to oppose cm the
other.
There appears to be no doubt that the resistance set up, if not
adequate for the destruction of the organism, even contributes
indirectly to an increase in the virulence of the latter. Thus, an
organism, e.g. t the Bacillus tuberculosis of reduced virulence in
milk, after inoculation was found to gain in virulence according as
the distance from the site of inoculation increased. The resistance
offered to the invading organism being insufficient to destroy it,
trained it, as it were, to develop more inimical powers of attack, so
that, as the bacilli overcame successively the barriers placed in their
path, they increased not only in virulence but in number also, just
as the Virus fixe of rabies was obtained by Pasteur by means of
passage of the street virus through a series of animals. This process
has been designated a 1 reinforcement by relays.* The overcoming
of this resistance by an organism of feeble aggressive powers of
course takes time whether the delay is caused by the small number
of organisms, or by their being of reduced virulence, or by the
greater degree of resistance set up leading to a 4 relatively reduced
virulence.* One or other of these, acting singly or in combination,
may be adduced to account for latency.
Where cases are met with in which death occurs at such a tender
age as 22 days (No. 128),* 24 days (No. 214), 29 days (No. 180),
and 7 weeks (Nos. 63 and 114), one is naturally led to enquire into
the question of congenital and hereditary tuberculosis.
The possible modes of congenital infection are, of course, either
by the ovum or spermatozoon, or via the placenta from a tuberculous
mother. As regards the first of these, as Adami states, ‘ the microbe
of an infectious disease cannot be a constituent of the biophore. At
most it can be an accidental inclusion in the surrounding non-
heritable matter of the cell.* The human ovum, being practically
• The serial numbers refer to those contained in a table showing the post-mortem findings
of the 300 cases of tuberculosis under consideration. The Editors regret that it is not possible
to reproduce this table.
2l6
free from yolk and not being phagocytic, cannot take up the bacilli,
while it is still more improbable that the minute spermatozoon could
carry them. Repeated observations go to show that the semen of
a phthisical patient (Rohlff, Gaertner) does not contain as many
as ten bacilli, and as the average human seminal ejaculation is said
to contain over 200 million spermatozoa, the chances that a single
spermatozoon which fertilises the ovum should carry a bacillus are
about one to twenty million. For practical purposes this mode of
infection may be regarded as impossible. The second method, via
the placenta of a tuberculous mother (or perhaps through the walls
of the foetal sac, or passing to the foetus before the sac forms), has
been shown to be possible; in fact, intra-uterine acquirement of
disease as distinct from inherited disease is proved. £uch trans¬
mission of the tubercle bacillus from parent to child in utcre, is
however, undoubtedly very rare. Thus Schluster, who examined
into the records of the reported cases, as mentioned by Latham, was
able to collect only twelve showing clinical evidence of tuberculosis
at birth. Furthermore, in the majority of instances, the organs of
foetuses bom of tuberculous mothers yield only negative results when
inoculated into guinea-pigs.
The theory of diathesis—the question as to whether the children
of tuberculous parents suffer from a greater liability to tuberculosis
from enhanced susceptibility resulting from 4 paratuberculous lesions *
due to toxic action on the germ cells—the hereditary transmission of
constitutional predisposition to tuberculosis, is being relegated more
and more to the realms of improbability as modem knowledge, with
its insistence on accuracy of detail, demands facts and proofs in
place of nebulous hypotheses.
The dominating factor in the incidence of tuberculosis is the
opportunity for infection; in other words, the greater the oppor¬
tunity for infection the greater is the incidence and the mortality
from this disease. Can we argue that the reverse of this also holds
good, namely, that a high incidence and mortality mean greater
opportunities for infection ? Probably we can, as will be shown
later when the local conditions are mentioned.
It would appear to be established that under certain circum¬
stances, no matter what the 4 diathesis ’ may be, infection can always
be effective. To prove that some children inherit a predisposition
217
to tuberculosis, the incidence of the disease amongst the offspring
of tuberculous parents and amongst equal numbers of non-tuberculous
parents would have to be compared, and, furthermore, it would be
essential that both should be subjected to the same environment
from birth, as McFadyean has pointed out. Even then we must
bear in mind the fact that the former will have been exposed to
infection during the intra-uterine period, so that the conditions for
comparison will not be strictly impartial. To compare, as has been
done, the death-rate from tuberculosis amongst members of the two
categories—the children of healthy parents on the one hand and
those of tuberculous parents on the other—is fraught with fallacy,
as the after conditions are so diverse. To make them more equable
it would be necessary for the children of healthy parents to be bom
in infected, houses or be placed there at birth, clearly an impossible
condition except by mere accident.
The children of tuberculous parents are thus handicapped in
several ways, each of which plays a part in rendering fallacious the
comparison of their death-rate with that among children of healthy
parents. The chief of these disturbing factors would be: (i) Intra¬
uterine exposure. (2) Birth in infected surroundings, and, therefore,
greater liability to infection while young. (3) The dose of infecting
material is likely to be large. (4) This dose is probably frequently
repeated.
That the intra-uterine period may be dangerous to the child
from the point of view of tuberculosis has been definitely proved,
though the cases on record are very exceptional. Thus SchmorT
and Birch-Hirschfeld, and Schmorl and Kockel, have described
cases in which the bacilli were found in the placentae and the
foetuses of tuberculous mothers. More recently Friedmann reported
the finding of two cases of tuberculous infection by the placental
route.
So much for the question of hereditary transmission of the
1 Bacillus tuberculosis. As regards environment no one now disputes
the importance of this factor in producing the disease, but though
comparison is difficult in human beings owing to the variety of
disturbing factors, as has already been stated, in animals we can
better arrange conditions to suit our purpose. The experiments of
Trudeau may be referred to in this connection. He inoculated
2l8
rabbits with tuberculosis and allowed some to run wild, whereas
others were kept in a damp, dark place. Most of the former
recovered, while the latter rapidly succumbed.
In cattle an analogous condition is found. As McFadyean
states: ‘ All breeds and strains of cattle are susceptible to tuber¬
culosis, and when the environment is the same the incidence of the
disease is the same in all breeds and strains.' He found that the
conditions under which the cattle are bred and reared constituted
the most important contributory factor according to the opportunity
supplied by the environment for the transmission of the bacilli.
Regardless of breed as a separate factor, the proportion of cases of
tuberculosis furnished by any breed is high or low according as the
cattle are or are not in close association with other diseased animals.
Calmette's experiments lend additional support to the same fact.
He found that cattle which had been once infected nearly always
recovered if they were carefully isolated, but if kept in prolonged
contact with tuberculous animals they themselves became actively
tuberculous.
So in man no race is exempt; in other words, there is a racial
susceptibility to tuberculous infection, but there is not sufficient
evidence at present to enable one- to say that any special
predisposition is handed on by tuberculous parents to their
offspring.
We may summarise the matter by saying, firstly, that hereditary
transmission of the bacillus is so rare that for practical purposes it
may be declared negligible; secondly, that the incidence of tuber¬
culosis depends on two main factors, namely, the dose received and
the virulence of the strain inoculated; in other words, the degree of
exposure to infection and the resistance which the inoculated subject
is able to put forwar^, this, in turn, being dependent largely on his
environment; thirdly, that there is not sufficient evidence, at present
at all events, to afford support to the theory that there is such a
thing as inherited predisposition to tuberculosis.
Before passing on to a more detailed description of some of the
300 cases of death from tuberculosis, the morbid anatomy of which
forms the basis of these studies, it will be advantageous Jo say a
few words on the extent to which the foregoing points are
exemplified in the conditions prevailing in Hongkong.
219
Generally speaking, the prevalence of the disease is closely
connected with social and economic conditions-^-overcrowding and
slums, poverty, insanitation, and squalor. The ingestion of
tuberculous milk, which is supposed to play a large part in the
production of tuberculosis, especially intestinal tuberculosis, in
children in England, has no influence here, for the Chinese children
do not drink milk, nor does it arise here from the use of tuberculous
meat.
The problems of tuberculosis in Hongkong are really social
problems, and are, therefore, intimately connected with those of
public health. The main causes of the prevalence of the scourge are
the predisposing ones of overcrowding of the poor and the fact that
the rooms inhabited by them are dark and the sunlight rarely enters
them. They are still further darkened by gratings and shutters.
With the first of these there is little if anything to be done; the
population is great and the space for their accommodation relatively
small; not only is floor space inadequate, but window space is less
than it should be, and the windows are often closed and made of
opaque or coloured glass, so that the penetration of light is reduced
to a minimum. This latter question of the darkening of the rooms
can only be improved by education.
In an interesting paper on * Sanitary Progress in Hongkong,”
Dr. W. W. Pearse, the Medical Officer of Health, states with
reference to the housing of the poorer Chinese that, the area being
limited and the Chinese population large and constantly increasing;
building sites have become very expensive so that the streets
inhabited by. this class of person are narrow, and the houses fronting
on them high; in some cases the height is five times the width of
the street.
' It is usual/ writes Dr. Pearse, 1 to speak of the Chinese houses
as being of one or more storeys, but as each storey was generally
let separately, and as the occupiers of each storey used it as they
would have used their single storey house in their native village in
China, in effect these Chinese houses in Victoria were piled one on
another, three, four and five high !
‘Another factor has contributed curiously to produce the ill-
designed Chinese house of Hongkong. The floors of these houses
are supported by China fir poles. A pole of more than fifteen feet
220
long of sufficient strength for a floor joist' is not readily procurable.
This has limited* the width of storeys to fifteen feet.
‘ In order to provide as much accommodation per floor as
possible, i.e. t to make building pay, the area of a floor has been
obtained by increasing its depth out of all proportion to its width.
‘ Hence a Chinese house in Hongkong has been a veritable tunnel
fifteen feet wide and forty, fifty and even sixty feet deep.
‘ Excepting comer houses only, windows to light and ventilate
these tunnels were possible only at the front and in front of these
were verandahs.
‘Windows at the rear of houses were not at first considered
necessary, and many houses were built back to back, with no yard
or ventilating shaft between them. On each floor the rear portion
was cut off by a partition wall to form a kitchen. Such kitchens
were small, generally under 150 square feet in area, very dark, and
often with no means of ventilation other than the door communi¬
cating with the main portion of the floor. These kitchens were
drained by vertical earthenware waste pipes which, in the case of
back to back houses, were carried down through the building to
discharge over a gully trap in fhe kitchen on the ground floor.’
From time to time laws have been passed to improve the housing
conditions by providing open spaces of a defined minimum area
at the rear for ventilation. Later, all new houses were required to
have larger back yards, and also scavenging lanes. Further,
regular systematic cleansing of the Chinese houses was instituted
(in 1903) and certain repairs effected. These measures were really
directed towards the eradication or limitation of plague, and were
aimed at affording protection from infestation with rats.
The method of cleansing the houses is carried out in the following
manner four times a year : the furniture is turned out, the bed-boards
and such like are dipped in 1 per cent, kerosene in water, in order to
get rid of vermin; the floors also are sprayed with the same liquid.
All rubbish and dirt generally are removed, and an inspector makes
a careful examination to see if any floors need repair, if any gratings
are missing, and so on.
The accompanying diagram, kindly drawn for me by Dr. W. J.
Woodman, late Acting Medical Officer of Health for. Hongkong,
and now M.O.H., Kewleen, shows the arrangement and mode of
221
occupation of a typical ‘ floor ’ of a modern Chinese house in these
districts.
The foregoing constitute the main predisposing causes, but the
direct cause of the prevalence of tuberculosis in Hongkong is,
facile princeps , the expectoration habit. Nearly all writers on
tuberculosis in the tropics note this peculiar trait. In the houses
of the poor a person will expectorate anywhere on the floor, or, if he
is ill in bed, on the bed clothes. Owing to the unavoidable over¬
crowding in the poorer districts the children play about on the
floor, putting everything into their mouths, breathing a vitiated
atmosphere and one often tubercle-laden. These are the reasons for
the extensive prevalence of tuberculosis among the native population
here and for the relatively high proportion of cases met with among
children. A subsidiary but analogous cause is that manner of
feeding their infants which many mothers, themselves tuberculous in
some instances at least, indulge in, of first chewing the food and
then placing it in the mouths of their babies.
Further, experiment has shown that there is considerable danger
of transmission of tubercle bacilli from one person to another by
means of eating utensils, if these are uncleaned. In China the eating
together in common, the using of the same general dish, the insertion
of individual chopsticks into the food supplied for the meal, the
transference of food with the chopsticks of one person to the mouth
or dish of another—all these afford a ready means of conveyance of
the organism, but readily preventable when it once comes to be
recognised.
These, however, are not so important as the one first named—the
222
expectoration habit—for intestinal tuberculosis, as shown in this
series, is not the common mode of infection of children out here.
Dr. Chausse compared the infection arising from dried sputum
and that from droplets of sputum and saliva directly inhaled, and
laid stress on the fact that it is exceptional for those living with
tuberculous subjects to receive directly the cough of the latter, and
the droplets are very soon diluted with a large quantity of air, and
thus danger is lessened. Further, these droplets dry very quickly
and their virulence is probably much diminished in ten days.
Deposition occurs in a few hours at most, and, owing to the
deposition and the drying, the droplets become reduced to the
condition of infective dust. ‘ Les gouttelettes bacillaires elles-meme,
comme facteurs de contagion de la phthisie jouent la plus grande
partie de leur role apres leur depdt et sous la forme de poussi£res
seches, de nouveau mobilis6es dans Tatmosph^re/
When no precautions are taken, and this is the rule, the clothes,
bedding, etc., are soiled by the virus in large quantity. Then the
making up of the bed, brushing of clothes, sweeping the floor,
distribute the organisms so that healthy contacts become exposed to
infection every day and all day long, for this dust may remain
suspended for several hours. Moreover, currents of air, movements
on the part of the patients, continually put the dust in motion.
Then, when the colder months come on the Chinese take out their
old clothing which has been stored away in some cupboard or dark
comer and wear that over their other garments. When the house
cleansing takes place these clothes are not disinfected, so that they
are nearly, if not quite, as infectious as when they were laid aside
at the termination of the previous cold weather.
Nuttall has estimated that a patient with open tuberculosis may
expectorate four thousand million bacilli daily. In a dark, shut-in
room the chances of infection for a child brought in contact with
such a case are enormous. During the crawling age these chances
are still greater. It will be seen on perusal of the age table appended
that of the two hundred and twenty-five cases under ten years of
age, one hundred and sixty-three, or 72 8 per cent., were three years
old or less, and one hundred and eighty-five, or 82 2 per cent.,
under four years. Further, Dieudonn6 of Warzburg (1903) has
proved that the nasal secretion and dirt on the hands of children
223
during the crawling age, say nine months to three and a half years,
in a very large proportion showed the presence of tubercle bacilli.
The opportunities for the repeated infection of a child of the crawling
age in contact with a person suffering from open tuberculosis, or even
living in a room previously occupied by such a person, are enormous.
The danger of infection from tuberculous members of a family is not,
according to the weight of evidence now available, due to the mere
‘ tendency to infection/ or even to the increased probability of
infection per se , but is to be ascribed to the probable large size and
frequency of the infecting doses. It has been shown experimentally
that small repeated infections may be protective, but large ones
overwhelm. Thus is explained the apparent antagonism of
Calmette’s and of Bryant’s experiments. The former have already
been mentioned with reference to single or repeated infection of
cattle, either diiectly or by keeping them in contact with tuberculous
animals. Bryant, after injecting guinea-pigs with a small number
(eight) of bacilli daily for a time and then every three days for a
period of four months, found lesions indicative of great resistance.
•The conditions m Hongkong, as has been pointed out, are
analogous to the experiments of Calmette. It is needless to labour
this point further, but in conclusion mention may be made of
Theobald Smith’s statement in the Journal of the American Medical
Association : * The resistance of the tuberculous animal to super¬
infection is readily broken down by slightly increased dosage, and
is successful only when very minute doses come into play.*
As already mentioned, the prevalence of tuberculosis is closely
connected with social and economic conditions—overcrowding and
slums, poverty, insanitation and squalor. With regard to over¬
crowding, little, if anything, can be done; the space available is
limited, and the population is relatively large and is increasing.
The question reduces itself, for practical purposes, to the finding of
a remedy for the spitting habit in the first place. The use of food
utensils, bowls, chopsticks, etc., in common probably plays a very
subordinate part, as has been stated above. To exact, or rather to
inflict fines for spitting in public would be useless for several reasons :
firstly, the offenders cannot afford to pay fines; secondly, the native
police are every bit as bad offenders in this respect as the rest;
thirdly, I am convinced that it is the expectoration on the floors of the
rooms in which the people live that is the chief source of the bacilli,
224
so that, even if expectorating in public out of doors were stopped—
1 a consummation devoutly to be wished’ on other accounts than the
risk of disseminating tubercle bacilli—there would be no deterrent
to a man exercising the privilege in his own rooms, unless his wife
objected, and this she would not do seeing that she also freely
avails herself of a Jike privilege. We are, therefore, reduced to the
slower but more certain method—Education, and the question
naturally fellows: How and by whom? European doctors would
be of little, if any, use directly. They do not come into contact
with the poor to any extent, and they would not be listened to even
if they could succeed in making ’ themselves understood. The
Chinese doctors would have a better chance, and, therefore, much
may be hoped from the locally qualified practitioner who comes more
into contact with the people, both at the hospital during his years
of training and also after he graduates and goes into practice.
Better still would it be to attempt to instruct the people in their
houses, and this, I venure to suggest, could be accomplished in the
following way :—The Y.M.C.A. and the Y.W.C.A., to which many
of the better educated classes belong, have courses of lectures from
time to time on various subjects. Like all the laity they are
interested in medical questions, and there is no reason why popular
(or, to use Huxley’s preferable term, people’s) lectures and
demonstrations should not be given at these institutions, and such
lectures cn medical subjects never fail to draw. This would form
a nucleus, and from among the members, the women more than the
men, some would certainly be found to spread the glad tidings of
the gospel of health, at first in their own homes among their servants,
and later tactfully to promulgate the doctrines they had been taught.
After a time a regular system of district visitors (in the medical sense)
might be inaugurated; these would not only instruct the mothers in
child welfare and the dangers of feeding their children in the way
indicated earlier, but would instil into the adults, and, through
them, the growing children health principles generally. The
teachers of the Chinese attending British schools might also with
advantage incorporate elementary hygienic principles as object
lessons for their pupils. Then the question of the establishment of a
tuberculosis dispensary would come up for consideration. The Chinese
are very fond of their children, and the appalling mortality from
tuberculosis must be ascribed to ignorance and not to perversity.
Showing distribution of the 300 cases according to age and sex.
Total
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REFERENCES
Adami, J. G. (1909). Principle* of Pathology. Vol. I, p. 184.
Burnet, Et. (1915). Ann airs de Vlustitut Pasteur. Vol. XXIX.
Chausse, P. * Nouvelle* Recherche* *ur la Contagion de la Tuberculo*e par l'air expir£ pendant
la toux.' Ann. de Vlnstitut Pasteur.
Delepine, S. * Contribution to the Study of delayed or “ latent ” tuberculous infection.*
Annales de Vlnstitut Pasteur.
Friedmann (1905). Vircboto’s Arcbiv . CLXXXI, p. 150.
Latham, A. (1908). Discussion on 1 Heredity and Disease.' Royal Society of Medicine.
Mcpadyean, J. (1908). Discussion on ‘ Heredity and Disease.' Royal Society of Medicine.
Nuttall (1911). Quoted by Cautley in Discussion on * Portals of Entry in Phthisis.' Proc.
Roy. Soc. Med.
Pearse, W. W. * Sanitary Progress in Hong Kong.'
Schmorl, and Birch-Hirschfeld (1891). Ziegler's Beitrage, . Vol. IX, p. 428.
-and Kockel (1894). Ziegler’s Beitrage. Vol. XVI, p. 313.
Smith, Theobald (1917). Journal of Amer. Med. Assoc. Vol. LXVIII.
THE PREVALENCE AND CHARACTER
OF TUBERCULOSIS IN HONGKONG
BY
HENRY HAROLD SCOTT
(Received for publication I ]uly % 1921)
II. THE PORTALS OF ENTITY AND MODE OF SPREAD
OF TUBERCULOSIS
The skin route as a portal of entry for tuberculosis which
becomes widespread is negligible; the nose and mouth are the only
ones worthy of serious consideration, and from the bacillary point
of view a certain amount of exchange between the respiratory and
alimentary tracts is possible; inhaled bacilli may be returned by
ciliary action of the epithelial cells and swallowed, while the latter
may in turn rise to the level of the larynx and be inhaled.
That bacilli when ingested can pass through the intestinal walls
and affect the mesenteric glands without leaving any trace of their
passage is a well-recognised fact, and they may also traverse or
grow in and be transmitted from these glands to the lungs and
elsewhere.
Experimental evidence goes to show conclusively that a very
much larger dose of bacilli is needed to set up the disease by way
of the alimentary tract than by the respiratory. In fact, there is
considerable difficulty in infecting animals by feeding. As Cobbett
states: ‘ The intestine is well guarded. It is otherwise with the
lungs. While not open to the attack of common bacteria in anything
like such numbers as is the intestine, it (i.e., the lung) has not
developed equal powers of defence, and tubercle bacilli when they
enter the bronchi can effect an entrance (i.*., penetration) and cause
tuberculosis in numbers which would be powerless for harm if
swallowed/ Thus, Gebhardt, working with diluted sputum, found
that eight hundred bacilli sufficed to infect by inhalation, while
ten to twenty millions failed when swallowed. Kossell, Weber and
Heuse found that 1 milligramme would infect calves t>y inhalation,
while one thousand times this dose given by feeding only caused
228
minimal lesions. More recently, Weber and Titze showed that it
requires at least io mgrms. of bacilli to infect a calf by feeding,
while i/ioo mgrm. may succeed when given by inhalation. Findel
infected dogs by inhalation with doses down to 014 mgrms., while
feeding with doses up to 63 mgrms. produced no effect whatever.
Also sixty-two bacilli would constantly infect guinea-pigs by
inhalation, even twenty sufficing in some cases, and in very young
animals even five. He gave other guinea-pigs doses up to twenty
thousand bacilli by feeding, but failed altogether to infect. Many
species of animals, including calves, goats, dogs and guinea-pigs,
are vastly more susceptible to infection by way of the air passages
than by way of the alimentary canal.
The same investigator, Findel, again showed that five million
tubercle bacilli when inhaled by a dog caused extensive tuberculosis
of the lungs, while one thousand two hundred and twenty times that
dose when given by the stomach produced no effect. As regards
guinea-pigs, in some experiments as small a number as twenty bacilli
were sufficient to cause pulmonary disease when inhaled, whereas the
animals were able to withstand three hundred and eighty-two
thousand times that dose when introduced into the alimentary canal.
Passing from these animal experiments to human cases, an
interesting and instructive example was brought forward by Cautley
in which one hundred and fifty-one children and two hundred and
nine adults were unconsciously subjected to feeding experiments
by consuming milk from tuberculous cows. Of these three hundred
and sixty individuals 4 only two of the children were affected, and
they merely had a mild adenitis. . . They had taken the milk
for a year and a year and a half respectively, and the milk was from
cows with virulent disease of the udder. Such cases imply that to
secure human infection by tuberculous milk the requirements are:
youthful age, a badly infected milk, and the prolonged ingestion of
such milk.’
Judging from this and the foregoing experimental work, we may
say that the probabilities, when one finds what is apparently a
primary lung infection, are that the condition has actually arisen by
inhalation and not by way of the digestive tract. A more detailed
consideration of the site of origin and the mode of spread in
individual cases is given later with illustrations from this series.
The tonsils have been regarded by some as the portal of entry
229
for the bacilli, and they quote tuberculosis of these glands in support
of their view. But as there is usually a focus in the lungs as well,
it is rarely possible to say whether the primary site of infection was
tonsil or lung, or whether both may not be secondary to another
portal. As Austin says: * Examples 6f apparently isolated tuber¬
culosis of the tonsils should be regarded with reserve, as the presence
of latent foci in the lungs can never be excluded.* Excised tonsils
of forty-five children (aged 2 \ to 15 years) examined for tuberculosis
by inoculation into guinea-pigs, by histological examination of
sections, by cultures and by smears, all proved negative except one
inoculation; we' can say, therefore, that infection of the tonsils is
uncommon alone. In fact, in most cases of tuberculosis of the tonsils
lesions are found elsewhere, especially in the cervical glands, and it
is rare to find tubercle bacilli in tonsils of children without clinical
evidence of tuberculosis.
‘By those who hold that pulmonary infection arises through the
cervical glands,* says Dr. Whipham, ‘various routes by which the
tubercle bacilli may gain access to the lung have been suggested:
first, that from the cervical glands they enter the lymph stream, and
by means of the lymphatic vessels enter the venous system, and so
are conveyed to the lungs; secondly, that their path is from the
cervical to the supraclavicular glands, and thence to the apex of the
lung; thirdly, that from the cervical they pass to the bronchial
glands, and thence are conveyed to the blood-stream and the
pulmonary tissues, though in this connection it must be remarked
that no communicating channels between the cervical and bronchial
glands have been shown to exist.*
The usual modes of entry and extension are, of course, the
following: —
(1) Mechanically with air, food, secretions, and so on, as for
example when inhaled bacilli lodge in some spot and produce a focal
lesion or are returned by the action of the ciliary epithelium, as has
been mentioned already; or a focus having formed, the material
bearing organisms finds its way into a bronchus, the bacilli pass out
and are swallowed or are drawn into another bronchus to set up the
condition afresh there.
(2) By direct extension through the tissues with caseation of the
parts first affected and involvement of the new tissues by contiguity.
(3) By way of the lymphatic channels which become invaded by
230
this contiguous extension. The bacilli thus pass to the next lymph-
node, and so on by the lymphatic vessels till the thoracic duct
conveys them to the left subclavian vein and the lymphatic mode of
spread then becomes
(4) Dispersion or distribution by way of the blood-stream;
thence they spread by the right side of the heart to the pulmonary
arteries, and so to the lungs generally. Or, by the extension of a
focus and erosion of the wall of a blood-vessel, they may pass by
way of the pulmonary veins to the left side of the heart and thus to
the whole systemic arterial distribution. Or yet again, by opening
into a smaller artery will invade the area supplied by that vessel.
Cobbett, in his summing up, expresses his opinion in these
words: 1 While I have no doubt that tuberculosis is frequently of
intestinal origin, especially in children, inhalation is the common
mode of infection, not only in phthisis, but in other forms of
tuberculosis, especially those in which the bronchial glands seem to
be the parts first affected/
In temperate climates isolated primary tuberculosis of the
intestines, rarely seen in adults, is not uncommon in children
(estimated at 25 per cent, according to some authorities); in tropical
countries, judging from my experience in the West Indies (Jamaica)
and the Far East (Hongkong), this route, or rather portal of
primary entry, is not proportionately so common. Moreover, in
temperate climates primary intestinal tuberculosis in children is
usually ascribed, if not always traced, to the swallowing of the bacilli
in quantities in infected milk. This is certainly not the case here.
It is probable that the intestinal infection is in the majority of
cases secondary to lung tuberculosis from swallowing the sputum,
even in children, and when definitely primary is due to direct
ingestion of the bacilli from dried sputum in infected dwellings.
With this preliminary clearing of the ground we are in a position
now to examine the extent to which these conditions as regards the
portals of entry are borne out in my series.
Of the whole three hundred cases there were two hundred and
nine in which the primary portal of entry appeared to be by the
respiratory tract, i.e ., 69*66 per cent., and in seven others which are
discussed later there was considerable evidence in support of the
* 3 *
same portal, bringing the total to two hundred and sixteen, or
72 per cent.
Only thirty-two have been definitely determined as being of
alimentary origin, i.e . 9 1066 per cent.; five more, discussed below,
were very likely alimentary; this would bring the total to 37, or
12*33 P eT cent.
In four other cases there was a possibility of almost simultaneous
entrance by way of the respiratory and alimentary tracts. In the
remaining cases the primary portal was uncertain. A few remarks
are given subsequently on each of these.
The number of cases occurring in adults in this series is
proportionately small, insufficient at present to form a useful basis
for study in detail, and the main objects of this investigation have
been the prevalence and character of the disease in children. Of the
three hundred cases there were two hundred and twenty-five of ages
up to ten years, and each consecutive hundred contained very similar
proportions; thus there were seventy-six in the first, seventy-three
in the second, and seventy-six in the third. As regards the sex and
ages of those comprising the three hundred, the table (see p. 225)
shows that one hundred and fifty-five were males and one hundred
and forty-five females. It will be noted that after adult life the
cases in males preponderate. Whereas in children up to the age
of ten years the proportion of males to females is about as 4 : 5, after
the age of twenty years the proportion is as 3*6 : 1. This does not
mean that the males are more subject to tuberculosis in adult life
than are females. When ill the Chinese exhibit strongly the homing
instinct, and make every effort to return to their birthplace to die.
The females can more easily do this; the males would if they could,
but they in many cases have to remain at work until it is too late.
Of the two hundred and twenty-five children under ten years
of age there were one hundred and twenty-six females ( i.e.,
56 per cent.) and ninety-nine males (44 per cent.). This is not of
much significance when we bear in mind that the female child in
China is not considered of much account, and among the poorer
classes from which these, records have been made, the females are less
well looked after. This is shown by the relative proportions of sexes
for the early months of life.
232
Of the two hundred and twenty-five cases under ten years of age,
then, one hundred and forty-eight, or 6577 per cent., were found
in whom the portal of entry was respiratory, and in five others
(discussed later) there was a strong probability of a respiratory
origin, in which case the percentage increases to 68.
Only thirty-one of these two hundred and twenty-five cases
appeared to be definitely of alimentary origin, i.e ., 1377 per cent.,
while among those of ‘ uncertain portal * were three others in whom
the evidence for the alimentary route had considerable weight; if
these be included the total percentage would be 15*11. Cases of
isolated primary tuberculosis of the intestine are very rare here in
my experience. In fact, only four were met with, namely, Nos. 7,
8, 81, and 112. The great majority, then, have a respiratory portal
of entry, the proportion to alimentary being as great as between
four and five to one.
In the following the primary portal was not determined with
certainty. It will be of interest to give a brief note on each of them.
No. 15. Female, aged 22 months.
The extensive spread of miliary tubercles in the lungs, the meninges, liver,
spleen, etc., indicate spread by the systemic blood-stream, possibly after the
pulmonary circuit, but the only focus found was the caseating right broncho¬
pulmonary gland ; how this became affected without some preceding lung focus
I cannot say, but minute search failed to find any. Except for the gland all the
tubercles appeared to be of the same age and recent.
No. 27. Female, aged 1 year.
There were two foci in the lower lobe of the right lung and miliary of the
same lung with corresponding glandular affection, the gland at the left hilus
probably by extension, the left lung not being affected. The intestine showed
tuberculous ulceration and the mesenteric glands were in large caseous masses ;
the spleen was the only viscus with tubercles. If the primary portal were alimentary
and the spread to the lungs via the thoracic duct, one would expect both lungs
to be involved. In favour of the respiratory origin of the lung condition are the
foci in the right lower lobe and the glandular involvement. The ages of the
pulmonary and the intestinal conditions would appear to be about the same, and
there is the possibility of the simultaneous entry by both routes.
No. 33. Female, aged 4 years.
A few miliary tubercles were scattered throughout the right lung and over
the right visceral pleura ; the superior tracheo-bronchial gland on the right was
enlarged and contained creamy pus; the hilus glands were swollen. The left
lung and glands showed no involvement. There were a few miliary tubercles
over the spleen surface ; miliary also at the base of the brain, especially about the
interpeduncular space. Minute search failed to find any focu’s, unless the glandular
abscess constituted it
*33
No. 34. Male, aged 1 year.
In this case the broncho-pulmonary glands and some of the tracheo-bronchial
were enlarged and caseous ; minute search, however, failed to find any focus
in either lung. The mesenteric* glands were also, some of them, enlarged and
caseating, but not adherent. The state of the mediastinal glands would warrant
one in expecting to find a lung focus. In this connection may be quoted the
results of experimental work by Calmette, Guerin, and Breton. They found
that in guinea-pigs dying in two to four weeks after being fed on the bacilli the
mesenteric glands (especially the superior deriving from the small intestine) were
enlarged and inflamed, although no trace of any intestinal lesion could be deter¬
mined. After 6-7 weeks these glands were caseous in greater or less degree and the
lungs showed involvement by miliary tubercles with affection of the corresponding
tracheo-bronchial glands. Hence in the present case the probability might seem
to be primarily alimentary setting up mesenteric adenitis and mediastinal glandular
affection secondary to this.
N. 35. Female, aged 2 years.
Infection extensive : broncho-pneumonic phthisis of the right lung with
cavitation in the middle lobe ; the left lung contained miliary tubercles all through,
but no focus; mediastinal glands, as would be expected, were more affected on
the right. The intestines showed numerous ulcers, one having perforated, and the
mesenteric glands were large and caseous; the liver and spleen showed miliary
tubercles, the latter more than the former, but the kidneys and brain showed
none. The intestinal ulceration being so extensive, together with the condition
of the mesenteric glands, favour the infection being primarily alimentary whence
extension took place via the thoracic duct to the lungs, though the right lung
was more affected than the left. On the other hand there had been time for
cavitation to arise so that the intestine might have become involved through
the swallowing of infected sputum. Here again the respiratory and intestinal
conditions did not appear to differ greatly in age, and it is quite possible that the
two-fold route was taken. One must always remember that to infer the relative
ages of two tuberculous lesions from the stages present is a proceeding liable to
fallacy.
No. 39. Male, aged 8 years.
This case showed widespread infection though the course of the extension
is difficult to trace. The oldest sites would appear to be : (i) The left mastoid
which was caseous and necrosed. As there was middle ear disease this may have
been a condition apart from the tuberculosis; although caseous and necrotic,
there was no actual proof of its being tuberculous in nature, (ii) Chain of large
caseous cervical glands* more on the right than on the left, (iii) Thick caseous
adherent peritoneum and mesentery as a vast cheesy sheet, (iv) Chain of large
caseating mediastinal glands in relation to both lungs, (v) Focus in upper lobe
of left lung, (vi) Mesenteric glands in large caseated masses. Other serous
membranes—pleura and meninges—showed merely small miliary tubercles, in the
latter limited to the base and not very numerous; the viscera—liver and spleen—
showed none on section, merely a few on the surface ; the kidneys none, and the
intestines none, except miliary on the peritoneal surface. The disease would,
therefore, not seem to have been a generalised blood-stream infection.
Was the primary portal of entry the intestine, affecting the mesenteric glands
and the peritoneum without leaving any trace in the intestine itself, and thence
spreading via the thoracic duct to the lungs ? Or was the mastoid primary, thence
2 34
affecting the cervical glands (but the mastoid affected was the left while the glands
were more marked on the right), and so by the blood-stream the lungs ? Or
were the cervical glands affected via the tonsils Xthey showed no signs themselves),
and thence the ear and lungs ? Or, lastly, was the primary route respiratory,
thence to the mediastinal glands and to the blood-stream, and by the sputum to
the alimentary canal ?
No. 50. Male, aged 3 years.
Focus in the left upper lobe and miliary throughout both lungs ; extensive
intestinal affection—ulcers in both large and small intestines, mesenteric glands
in caseous masses. Miliary tubercles in liver, spleen, and kidneys, but not in the
meninges or brain. The alimentary appears a little older than the respiratory.
The question is whether there was infection by double route, respiratory from the
focus in the upper lobe of the left lung, and the intestine separately, or whether
intestines first, thence by the mesenteric glands to the thoracic duct and lungs
and then generalised ; but, if so, why should the meninges not show any involve¬
ment ? Or, again, the lung focus may have been the primary condition, the
intestines have become affected from that by swallowing the sputum, and from
the intestines the mesenteric glands, the lymphatic system, thoracic duct, superior
vena cava, and again the lungs to produce the miliary condition there.
N. 52. Female, aged 4. years.
The distribution in this case was fairly general but showed certain peculiarities.
There were a few minute tuberculous ulcers in an early stage in the small intestine
with tubercles on the corresponding serous surfaces. The mesenteric glands,
though not large nor adherent, were caseous. The peritoneum and omentum
were thick and studded all over with tubercles varying from miliary to the size
of large hempseed. As regards the viscera, the liver and still more the spleen
showed miliary and grey tubercles, but on the surface only, none were seen on
section. A remarkable thing, however, was that the ovaries were both enlarged,
almost to the size of adult ovaries, and contained caseous masses, or rather consisted
of a caseous mass in each case, the left a little larger than the right. The right
lung contained two foci in the lowest lobe, the size of a pea, miliary tubercles
throughout, and a more closely-packed mass of grey tubercles subapical in position ;
the superior tracheo-bronchial and tracheal glands were enlarged and caseous.
The left lung showed miliary tubercles throughout, with closely-packed milia on
the somatic pleura ; none such were seen on the right. The pericardium also
showed several miliary tubercles. There were a few scattered tubercles along and
just above the right Sylvian fissure and some at the interpeduncular space. Which
was primary is largely a matter of conjecture. For a single organ the ovary was
the most extensively involved, each being converted practically into a cheesy,
almost creamy, mass. The two lung fod had a fairly hard caseous consistence
and were well defined. The alimentary tract tuberculosis and the serous mem¬
brane involvement, particularly the latter, were extensive, of the miliary type,
though, as stated, a few of the glands were caseous.
No. 54. Female, aged 7 months.
Distribution widespread; from appearances the respiratory and alimentary
involvements seemed to be of about the same age. There was a focus at the lower
edge of the right upper lobe, extending a little into the middle lobe; miliary
and grey tubercles were present elsewhere in the lungs, and the glands were caseous.
Alimentary : ulcers in the small intestine, mesenteric glands adherent and caseous,
2 35
but not fused ; liver and spleen showed a few miliary tubercles. The right kidney
had a small mass occupying the pyramid at the lower pole. The meninges at the
base contained several miliary tubercles. A cervical gland on the right side was
enlarged and beginning to break down in the interior. The question is whether
the lung focus was the primary (the enlarged gland on the same side would support
this) from whence extension occurred to the blood-stream to be generalised and
by swallowing of the sputum the intestinal condition ; or, whether from the lung
primarily to the intestine and mesenteric glands, and by the lymphatic system
to the pulmonary circulation to set up the miliary tubercles there ; or, lastly,
lungs and alimentary tracts together, then generalisation from the lungs to the
systemic vessels and from the alimentary tract by the glands to generalisation in
the lungs again.
No. 55. Male, aged 3 years.
The right lung showed tuberculous broncho-pneumonia of the middle and
lower lobes; the upper appeared to be quite unaffected. The middle contained
at the lower part a cavity the size of a cobnut. The left lung consisted of three
lobes and showed small ulcerated cavities throughout, the average the size of a
pea, but rather larger in the middle lobe. Against the lung being primary is the
fact that in spite of the extensive tuberculous condition of the organ the glands,
though a little swollen and congested, showed no obvious tuberculous infection.
On the other hand, the mesenteric glands were enlarged and three of them were
caseous, almost chalky, though the intestines showed no change to the naked eye.
The possibility is that the alimentary route was primary and thence affecting the
lungs by the lymphatic system, the upper lobe escaping, and the hilus and other
mediastinal glands not having become tuberculous before death supervened; or
that, since the pulmonary condition was broncho-pneumonic (rather than miliary
by way of the pulmonary circulation), the disease had originated by the respiratory
route and the alimentary tract had become involved by swallowing the sputum.
No. 58. Male, aged 14 months.
In spite of the age of this child the conditions were such that the general
distribution of the disease makes it a matter of conjecture as to the primary portal
of entry. There were signs of tuberculous broncho-pneumonia throughout both
lungs, with ulceration in the left lower lobe. Generalisation by way of the blood¬
stream may have taken place from this, causing the deposition of tubercles in
liver, spleen, kidneys, and brain. The intestinal ulceration and the masses of
caseous mesenteric glands may have arisen from swallowed sputum, but the intestinal
condition appeared to be equally advanced as the pulmonary and may have arisen
independently. The pericardium also was studded with miliary to pin-head
tubercles.
No. 59. Female, aged 4 years.
There was marked kyphosis from extensive involvement of the vertebrae.
The spinal site was probably the oldest. From the number of ulcers in the
intestines, the large and small both being involved, and the mesenteric glands
being in large caseous masses, the alimentary tract would appear to have been
attacked prior to the lungs. The latter showed miliary tubercles throughout,
but there was also a fair-sized focus (cobnut) just below the apex of the right
lower lobe in an early stage of caseation.
236
No. 65. Female, aged 8 months.
The peculiar limitation of the disease in the lungs to the middle lobe of the
right, which contained a distinct focus the size of a pea and small miliary and grey
tubercles throughout that lobe, would be in favour of a primary (or at least
independent) lung involvement, the alimentary tract being affected secondarily
from swallowing the sputum. The mesenteric glands, however, were in large
caseating masses and appeared to be of older standing than the pulmonary
condition.
No. 66. Female, aged 20 months.
In some respects similar to the last. In this case the lung affection was limited
to the right upper lobe in which there was a subapical focus, and in addition
generalisation of small tubercles throughout that lobe. The disease was much
more advanced in the alimentary tract—intestinal ulceration and large caseous
masses of mesenteric glands—and this would point to an alimentary portal of
entry. It is not improbable that the lung infection, though later, was independent
of the intestinal. The condition of the brain (cerebellar mass) and meninges
(miliary to hempseed on vertex and base) point to dissemination by the blood¬
stream, but no other viscera showed involvement, to the naked eye at least.
No. 68. Male, aged 3 years.
This case is remarkable in that, except for a hilus gland and a paratracheal
gland on the right side as large as a cobnut and filbert respectively and caseous,
no focus was found after careful search. There were sparse miliary tubercles
throughout both lungs: in the meninges they were numerous, especially at the
base, the interpeduncular space, the ependyma, along the Sylvian fissures, and a
few on the vertex. The spleen showed also a few milia on the surface. There
was no evidence of tuberculous affection of the intestines.
No. 70. Female, aged 7 years.
In this child there was a mass of caseating glands on both sides, involving
the inferior and superior tracheo-bronchial and the paratracheal, more on the right
than the left; the broncho-pulmonary were not apparently affected. No focus
was found in the lungs. The tonsils were not affected, nor the cervical glands,
and the spine was examined without success. The alimentary tract was free.
In the kidneys, however, there were in the left miliary and grey tubercles, not
numerous; in the right more, and arranged almost focally as a group at the base
of a pyramid towards the lower pole.
No. 92. Male, aged 10 years.
The primary portal in this case was probably alimentary. There were several
ulcers, more in the large than small intestine, however, and a thickened tuberculous
mass like a collar at the ileo-caecal valve. Mesenteric glands were enlarged, in
masses, and caseating. The lungs showed tubercles varying in size from pin¬
point to small pea throughout, but the mediastinal glands did not show any
tubercles visible to the naked eye, and only one was a little congested and swollen.
Neither brain, liver, nor spleen showed any signs, but the kidneys were severely
diseased. The right had the pyramidal areas hollowed and caseous; they had
run together and were lined by thick tuberculous matter. The left was in a
similar condition but to a less degree ; calyces opening into each other with a
lining of tuberculous pus. The tuberculosis was, therefore, widespread, but if
by way of the blood-stream it is peculiar that the kidneys should have been
237
extensively affected while the brain, liver, and spleen escaped. The lung condition
appeared to be more recent than the intestinal and may have arisen by way of the
pulmonary circulation as previously pointed out.
No. 95. Female, aged 8 years.
In this case one of the inferior tracheo-bronchial glands was caseous and one
of the superior in an earlier stage, both on the right side, but minute search failed
to find any tubercles in the lungs. The intestines were not affected, and no focus
was detected anywhere. The meninges showed miliary tubercles in fair numbers
at the base, fewer along the Sylvian fissures, and only an occasional one at the
convexity. No tonsillar or retro-pharyngeal affection was detected. If the
meningeal condition arose by inhalation, via the ethmoidal canals (as meningococcus
may pass), then we have no explanation of the mediastinal gland affection. Anyway
these do not appear to be accounted for, though the spread may have been from
them by the blood to the meninges.
No. 96. Female, aged 9 years.
In this case both the respiratory and alimentary tracts showed such extensive
involvement that the only doubt is whether the lungs, which were in a state of
tuberculous broncho-pneumonia throughout, with cavitation in the right upper
lobe and considerable ulceration in the middle lobe, were the primary seat and
the intestine was involved later through swallowing the sputum, or whether the
aljjnentary tract formed the primary portal of entry. The small intestine contained
tuberculous ulcers; the mesenteric glands were in large caseated masses and
adherent, and the peritoneum was studded with tubercles. Of course, there
may have been infection by both routes nearly simultaneously.
No. 100. Male, aged 5 years.
In this child there was a cavity in the right upper lobe the size of a medlar,
with grey tubercles surrounding it and smaller ones at a greater distance. No
other part of the right lung was affected and the left showed nothing. There
were pleural adhesions at the apex and direct communication with a tuberculous
ulcer the size of a crown-piec* above the right clavicle. Whether a supraclavicular
gland had become infected from the lung direct and had then opened on the
surface, or whether there was a primary affection of the skin which ulcerated and
extended to the lung is doubtful. There was a tuberculous skin affection with
early ulceration on the right cheek also. No other viscera showed any involvement,
and no cervical glands were found enlarged. This may be an example of the
mode of spread already mentioned.
No. 104. Male, aged 16 months.
In this the disease was widespread and the primary portal and mode of extension
• doubtful. The left lung showed two foci, one subapical in the lower lobe, the other
nearer the lower edge, and each was surrounded by a zone of small caseating
tubercles. The right upper and lower lobes contained miliary tubercles; the
glands, hilus, and tracheo-bronchial were caseous and enlarged. In the alimentary
tract there was nothing, but one or two mesenteric glands contained small foci.
Milia were present in the liver, more and larger in the spleen. The meninges
had masses of miliary tubercles at the base, and along the Sylvian fissures and in
the interpeduncular space; a few on the vertex and internal surfaces of the
cerebral hemispheres. The spine showed a large abscess the size of a Tangerine
orange, involving the nth and 12th dorsal and the 1st lumbar vertebrae. Sug-
238
gested method of spread : Lung (left), then by inhalation the right, by the blood¬
stream to the liver, spleen, spine (injury possibly determining this site), and also
by the sputum to the intestine and mesenteric glands; or, from the left lung
foci by the sputum to the intestines and mesenteric glands, thence by the lymphatic
system to the heart and pulmonary circulation, causing the miliary tuberculosis
of the right lung, and also by way of the blood-stream from the lung to the liver,
spleen, and meninges. But the spine focus must have been of considerable standing,
for there was marked caries and an abscess formed containing caseous matter.
No. 116. Female, aged 10 months. #
The right lung showed two distinct fod in the upper lobe : the larger subapical
the size of a cherry and caseous, the smaller the size of a pea, near the interlobar
fissure; the rest of the lung contained miliary tuberdes. The left lung showed
miliary to pin-head tubercles, sparse in the upper lobe, more in the lower. The
mediastinal glands were enlarged, with caseating fod. The mesenteric glands
(the upper set) were in a similar condition. The question is whether both the
respiratory and alimentary tracts were involved simultaneously, or whether the
lung focus was primary, the alimentary secondary, passing to the glands without
leaving signs in the intestine itself. Thence by the lymphatic to the pulmonary
circulation to set up the miliary tuberculosis there, and from the lungs via the
blood-stream to the meninges, liver, and spleen. But, if so generalised, one would
hardly expect the kidneys to escape altogether.
No. 118. Female, aged 3 years.
In this case the disease was exceedingly widespread and the portal of entry
is a matter of conjecture. There were tuberculous ulcers of the right cheek,
jaw, and neck; the right parotid, the submaxillary and cervical gkands on both
sides, were caseous. There was tuberculous broncho-pneumonia throughout
both lungs, the pleura showed many tubercles at the upper part of the somatic
layer and all over the visceral; the pericardium was also affected. The peritoneum
was studded with tubercles, miliary to pea, and the intestines contained a few
tuberculous ulcers, both in the large and the small; the mesenteric glands, both
upper and lower series, were enlarged, adherent, and caseated. The liver showed
a few miliary, the spleen more and somewhat larger. In the right kidney was a
small mass of aggregated tubercles at the lower pole, while in a corresponding
situation in the left was a definite caseous focus the size of a cherry. There were
aggregated miliary tubercles on the basal meninges, a few also on the vertical.
No. 137. Male, aged 10 months.
Disease was widespread and severe in this child, especially when one considers
the age. The lungs showed a cavity in the lower lobe, a caseous focus in 'the
middle, and miliary tubercles throughout the remainder of the right; in the
left there were two caseous fod the size of a pea in the upper lobe, a caseous mass m
as large as a walnut in the lower, and grey tubercles through the rest of the lung.
All groups of the mediastinal glands were involved. The mesenteric were enlarged,
caseous, and matted, but appeared to be more recent than the thoracic. There
were two tuberculous ulcers in an early stage in the small intestine. The femoral
glands on each side were the size of a small walnut and caseous (the origin of these
could not be discovered). The cervical glands on each side were caseous, especially
on the right. The right kidney contained a small aggregation of tubercles at the
lower pole ; the liver a few miliary tubercles, the spleen more and larger. The
meninges were not affected. As the respiratory and alimentary tracts were both
239
involved, and also some glands unconnected with either, it is difficult to say which
was the primary portal, but the lungs would seem to be most involved and of
oldest standing.
No. 141. Male, aged 10 years.
The oldest site in this case was to the right of the sixth cervical vertebra.
The bone was carious, and there was a sac with pus discharging from it along the
spine into the right pleural cavity (it may possibly have started as a retropharyngeal
abscess). The right lung showed miliary tubercles throughout, not very numerous,
but more in the upper lobe than in the other two. The left lung had none.
There was a caseous gland at the right hilus. Tuberculous sinuses were present
in both femora and in the left tibia..
No. 145. Female, aged i\ years.
Judging from the appearance, degree of caseation, etc., the primary portal
may have been alimentary, but the disease was very widespread, the brain showing
several foci. There were tuberculous ulcers in both large and small intestines,
and the mesenteric glands were in adherent, caseous masses. The cervical gland
infection was of fairly long standing also, those on the right containing creamy pus,
those on the left being in a caseated stage. Both lungs showed grey tubercles,
and the mediastinal glands were in caseous masses at the hilus and along both
sides of the trachea. The brain showed: three distinct foci in the left lobe of
the cerebellum, another in the right hippocampus, another in the left superior
frontal, all these about the size of a pea or a little larger, and in the right supra¬
marginal gyrus one as large as a cherry—an extensive series of brain foci.
No. 149. Female, aged 3 years.
It is possible that the alimentary portal was the primary as this tract showed
the greatest involvement (not a very safe indication, it is true). There were
numerous ulcers in both large and small intestines, the mesenteric glands, both
upper and lower series, were enlarged, adherent, and caseous. The lungs contained
closely aggregated miliary and grey tubercles throughout, but no focus, although
the mediastinal glands on both sides were caseous and the paratracheal on the
right contained creamy pus. The brain was considerably involved: there were
a few miliary tubercles in the basal meninges, many along each Sylvian fissure,
and a few on the opposed surfaces of the cerebral hemispheres ; in the cerebellum
at the fore part of the left hemisphere on the upper surface there was a ‘ solitary ’
tubercle, as large as a cherry, and towards the posterior another focus the size
of a haricot; on the under surface of the right hemisphere was one the size of a
filbert situated anteriorly, and at the posterior extremity another the size of a pea.
As regards the course, it does not appear likely that the large tubercles in the
cerebellum would be of the same standing as the miliary in the meninges. The
intestinal appeared to be the oldest, the lungs probably secondary to this via the
pulmonary circulation, for the distribution was equal over both lungs which were
granular everywhere from miliary and grey tubercles. General blood infection
was supported by the presence of a few tubercles in the liver, spleen, kidney, and
meninges, and probably an earlier blood infection set up the various brain tubercles.
No. 157. Male, aged 5 years.
Lungs were granular in appearance from the presence of numerous grey
tubercles. A gland at the right hilus was swollen and congested, but showed no
signs of tuberculosis in it. Intestines nil. One mesenteric gland, the size of a pea,
240.
contained small grey points. Both kidneys showed miliary tubercles, the left a
little more than the right. There were milia also in the liver, spleen, and
meninges. No focus could be found. The universality and evenness of the
pulmonary distribution point to a haemic infection of the lungs rather than a
respiratory.
No. 159. Female, aged 8 years.
The condition in the lungs and in the abdomen appeared to be of about
equal development. There were two foci in the left lung, one in the upper lobe
the size of a small cherry, caseous with a fibrous border, another in the lower lobe
slightly larger. The right upper lobe showed tuberculous broncho-pneumonia,
the middle and lower many grey tubercles. The mediastinal glands on both sides
were enlarged and caseous. The alimentary tract showed several tuberculous
ulcers in the small intestine, and the corresponding mesenteric glands were in caseous
adherent masses. There is apparently equal evidence for respiratory and alimentary
portals, and the stage in each is about the same. Possibly the double route was
taken nearly simultaneously.
No. 160. Male, aged 3 years.
Miliary tuberculosis of the lungs and meninges, a few also in the spleen. Two
paratracheal glands were swollen and contained two small caseous foci, and one
at the hilus was enlarged and caseous. No focus was found anywhere.
No. 164. Female, aged 6 years.
The difficulty in this case is to decide, in view of the widespread condition,
which was the primary portal of entry. There were old-standing ulcers beneath
each ear, with caseated and breaking-down cervical glands. The right lung
showed several foci, certainly not of recent production, one the size of a pea, deep
in the upper lobe, another similar at the base of the lowest; a small cavity in the
middle, and, at the upper part of the lowest, a large focus the size of a cherry,
caseous, with surrounding tubercles and early ulceration. In addition to these
there were milia throughout the lungs. There were tuberculous ulcers in the
intestines ; the mesenteric glands were enlarged, caseous, adherent. The meninges
showed numerous tubercles at the base and along the Sylvian fissures, more again
below the cerebellum. The vertex and inter-hemispherical aspects showed
tubercles in fair numbers. In spite of the lung condition the mediastinal glands,
though containing tuberculous foci, were not nearly as much affected as the
mesenteric. The respiratory and alimentary appeared to be of nearly equal age,
as regards the foci; the miliary state of the lungs may be a secondary result of the
intestinal. Doubtful whether : (i) Focal pulmonary, then intestinal, mesenteric,
generalisation in lungs and thence the brain ; or (ii) Primarily skin, then glands,
blood-stream and generalisation; or (iii) Skin, lungs, intestines (the last two
independently and simultaneously, as regards the focal state), and subsequent
generalisation.
No. 178. Female, aged 10 years.
Miliary tuberculosis of both lungs, fairly numerous and evenly distributed,
but rather more in the right lung. A gland at the right hilus and one paratracheal
caseous. Nothing in the alimentary tract. A few miliary in the meninges,
more at the base, but some also at the vertex. No focus detected anywhere.
2 4 I
No. 181. Female, aged 4 years.
Meninges showed general involvement, more aggregated at the base, in the
interpeduncular space, along the Sylvian fissures, and below the cerebellum ; few
on the convexity. The lungs showed widespread miliary tubercles, more numerous
in the right than the left. Mediastinal glands on the right were enlarged and
caseous, but no focus was found.
No. 203. Male, aged 5 years.
Miliary tubercles scattered in moderate profusion and evenly through both
lungs, but the source of these was not detected. The alimentary tract did not
show any involvement, but one mesenteric gland was the size of a pea, and on section
contained a minute caseous ’point. There was extensive tuberculous meningitis;
at the base they were closely aggregated and pus was forming; there were
also many along the Sylvian fissures and only a little less on the convexity. The
ventricles contained more than the normal amount of fluid. No focus was found.
No. 206. Male, aged 16 months.
Grey tubercles in the upper lobe of each lung and less in the right middle
lobe. The spleen showed a considerable number of miliary tubercles. The
meninges were extensively affected ; there was pus at the base with very numerous
tubercles, also along the Sylvian fissures, in the velum interpositum and below the
cerebellum. No focus was found, the other viscera yielding no evidence.
No. 210. Male, aged 5 years.
Here again no focus was found. The right lung contained miliary tubercles
in the middle and lower lobes, and a gland at the hilus was caseated. The upper
lobe of the left lung also contained miliary tubercles ; there was no affection of
the glands on the left; a few miliary were seen also in the spleen, while the
meninges were extensively involved ; tubercles were numerous at the base and along
the Sylvian fissures, and there were one or two on the convexity and on the opposed
surfaces of the cerebral hemispheres.
No. 211. Female, aged 5 years.
Apparently a haematogenous infection, but the primary focus was not found.
The lesions were miliary tuberculosis of both lungs, the spleen, and the basal
meninges.
No. 2ij. Female, aged 13 months.
The disease in this child was extensive and the decision as to the primary
portal of entry a matter of opinion. There was a caseous focus as large as a filbert
in the right lung, and grey tubercles throughout both lungs, while all groups of
the mediastinal glands were enlarged and caseous. The cervical glands were a
little affected. The mesenteric were in adherent caseous groups of about the
same degree of involvement as the mediastinal; there were miliary tubercles of
the liver and spleen, and a terminal meningitis. The mesentery and peritoneum
generally were studded with tubercles. From the state of the glands in the
thorax and abdomen there is no evidence for priority. The lung condition with
a caseous focus of such a size must be of considerable standing, whereas the intestine
itself showed one small ulcer only. On the other hand, the extensive affection
of the peritoneum indicated that the abdominal condition was not very recent.
On the whole, unless we regard the respiratory and alimentary as about coequal
242
in age, the most satisfactory explanation would be, the lung focus primarily,
alimentary secondarily, thence the peritoneum and mesenteric glands, and via
the thoracic duct to the pulmonary circulation and so setting up generalisation
there, and via the blood-stream (systemic) to liver, spleen, and meninges.
No. 222. Female, aged 18 months.
Though the mediastinal glands on the right side were enlarged and caseous,
no true focus was present in the lungs. The middle and lower lobes showed
grey tubercles of the minute bronchioles, while the right upper and both lobes
of the left contained miliary tubercles by the blood-stream. Some of the mesen¬
teric glands were the size of a small pea and had caseous points. The liver, spleen,
and ladney contained tubercles varying from small miliary to hempseed, and th6
base of the meninges showed them also. On the whole the sequence was probably :
Respiratory first, causing the broncho-pneumonia of the lower lobes of the right
lung, then alimentary to the mesenteric glands, without leaving signs in the
intestine itself; thence by the lymphatics to the pulmonary circulation and so
miliary generalisation in the lungs, and from there to the liver, spleen, kidney,
and meninges.
No. 223. Female, aged 4 years.
Extensive tuberculous meningitis, but no focus detected anywhere.
No. 227. Female, aged 3 years.
Lungs extensively affected with miliary and small grey tubercles, the right
rather more than the left. There were numerous tubercles in the meninges, and
a few in the liver and spleen, but no definite focus or indication of the primary
portal of entry.
No. 230. Male, aged 3 years.
Tuberculous meningitis most severe, but the origin of this could not be
traced. No focus was found, but there was miliary tuberculosis of the lungs,
liver and spleen.
No. 239. Female, aged 3 years.
In this case the alimentary, as evidenced by the state of the mesenteric glands,
would appear to be the oldest; on the other hand the caseous focus in the left
lung would be of older standing than the miliary and grey tubercles scattered
generally throughout the lung. Also the mediastinal gland corresponding to
the lung focus was completely caseous, like the mesenteric glands. The cerebellar
focus was not very recent, less so than the miliary in the lungs and meninges.
There are three possibilities: (i) Primary lung focus, alimentary secondarily,
then generalisation in the lungs and from there to the meninges (if so, whence
would the 4 solitary * tubercle in the cerebellum arise, unless by separate earlier
metastasis from the lung focus ?). (u) Primary intestinal, secondary focus in
lung and spread from there by inhalation to other parts of the lung, and by blood¬
stream to the cerebellum and meninges (but the cerebellar is a caseous node,
while the meningeal affection is miliary and apparently recent), (iii) Lung and
alimentary routes about the same time (judging from the glandular infection
those in the thorax and those in the abdomen appear to be about the same stage),
then from the lung to the cerebellum (focus), and from the mesenteric glands
to the right side of the heart for the production of the miliary and grey tubercles
in the lungs.
2 43
No. 275. Female, aged 6 years.
The lesions found were a general miliary infection of the right lung, and a
few milia in the upper lobe of the left; an enlarged and caseated paratracheal
gland on the right; sparse miliary tubercles in the spleen, and fairly thickly-
studded miliary infection of the meninges, more aggregated at the base. No
focus was discoverable, unless the mediastinal gland be regarded as such, but no
evidence of the communication of this with a pulmonary vessel could be made
out to account for the distribution in the lung, and there was certainly no focus
in the lung to which this gland could be attributed.
No. 283. Male, aged 4 years.
The lungs in this case presented miliary tubercles, rather more in the right
than the left, but not very numerous in either ; the hilus on the right showed
a large caseous gland, and one paratracheal on each side was caseated, that on
the right being the larger. No focus was found in the lungs to account for these
glands. There were a few miliary tubercles in the liver, more in the spleen.
The meninges were severely affected, at the base and over the right hemisphere.
In the latter, just below the cortex of the angular gyrus, was a mass of tubercles
focally arranged, aggregating to the size of a large pea or small marble. The
cerebral condition had probably arisen by vascular spread from the mediastinal
gland, but the source of the latter was not found.
No. 291. Female, aged 4 years.
The only focus detected in this case was a caseous mass occupying nearly
an entire pyramid at the lower part of the left kidney. There was miliary tuber¬
culosis of the lungs, but no focus. No infection of the intestines; only one
mesenteric gland showed anything and this was not so large as a haricot, but had
a small caseous point. There were also a few milia in the liver. The kidney
lesion was to all appearances the oldest, but whence this arose could not be
discovered.
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Calmette, Gu£rin, and Breton (1907). Ann . de VInst. Pasteur. Vol. XXI, No. 6.
Cautlry (1911). Discussion on 4 Portals of Entry in Phthisis/ Proc. Roy. Soc. Med.
CoBBirr, L. (1911). Discussion on ‘ Portals of Entry in Phthisis/ Proc. Roy. Soc. Med.
Findel. Zeitscbr.f. Hyg. Vol. LVII, p. 83.
Kossell, Weber, and Heuss (1905). Tuberkulose-Arbeiten aus dem kaiserlichen Gesundheit-
samte, Berlin. 5th Heft.
Weber and Titee (1910). Tuberkulose-Arbeiten aus dem kaiserlichen Gesundheitsamte,
Berlin. 10th Heft.
Whipham, T. R. (1911). Discussion on the * Portals of Entry in Phthisis/ Proc. Roy. Soc. Med.
245
MULTIPLE ANEURYSMS IN A CHILD
BY
R. H. KENNAN, M.D.
(Received for publication 3 January , 1921)
The patient is a well-grown, intelligent, active but anaemic boy
of four years old; the son of observant, healthy white parents,
American Missionaries on leave from Sierra Leone. The boy had
had about six attacks of fever, each of short duration, in Freetown,
and the latest of these occurred in October last. His tongue was
clean and moist and without tremor; his gums healthy; his pulse
regular and of good quality, with frequency of ninety-two beats per
minute while he was in the erect posture. His heart was free from
murmur and normal, and no enlargement of his liver or spleen or
lymph glands could be detected. There was no sign of rickets and
his urine (acid, 1,022) was free from albumen and sugar. His
parents suggested that there was oedema of his eyelids, but at the
time of examination (about 4 p.m.) I could not satisfy myself that
oedema was present.
The upper end of the left hypothenar eminence was slightly
enlarged, and prominent, and was the site of a pulsating tumour
which could be ' emptied * by pressure over it, and which then filled
again 4 per saltum.’ Pressure over the ulnar artery greatly
diminished, but did not entirely obliterate pulsation in the tumour.
The surface tissues over the tumour were unaltered in colour and
texture. Just above this tumour, and separated from it by a narrow
surface depression, was a second one of similar character.
There was no history of injury antecedent to the first hypothenar
tumour, which was first noticed in March, 1920; the second
swelling appeared in October, 1920: the parents state that it came
quickly two days after a fall, which, however, did not cause bruising
or damage to the skin. Subsequently a small pulsating little vessel,
slightly bluish in colour was noticed about the middle of the right
246
side of his neck, and some time after this, while his mother was
examining his neck, she found another larger pulsating, uniformly
oval swelling a little lower down, occupying the greater portion of
the base of the anterior triangle of his neck above the clavicle. The
skin over it was freely moveable, as was also the tumour from side
to side, but it was not capable of displacement vertically. Vigorous
pulsations were easily observable by sight and touch in the little
vessel; the pulsation's could be stopped by slight pressure below the
point where it lay immediately beneath the skin. The larger, lower
tumour could not be emptied by pressure over it, but the pulsations,
synchronizing with the ventricular systole, appeared to be distensile
in character, and with a stethoscope a loud bruit was audible over
it. The radial pulses were of equal volume, and no difference in
colour or temperature of the hands could be detected. It could not
be decided whether the wrist and hypothenar tumours were
separated by normal vessel or whether the surface depression was
due merely to compression by fascial bands. None of the tumours
caused any pain, and only when the larger neck tumour was firmly
pressed in an effort to empty it did the patient show any sign of
distress.
Examination of the blood showed Plasmodium falciparum infec¬
tion and also eosinophilia (about 12 per cent.). Several very careful
and prolonged examinations of the faeces yielded negative results.
There appears to be no doubt that the two distal tumours are
arterial aneurysms, and that both the abnormal conditions in his
neck are of the same kind. The alternatives of ‘venous pulsation,*
and ‘ enlarged lymph gland with transmitted pulsation * were borne
in mind at the time of the examination.
Macfie and Ingram (1920) have described ‘ Three cases of Cardiac
Aneurysms in Native Boys of the Gold Coast,* and discuss the
question of aetiology, and the suggestion is made that malaria may
have been a factor in causation of the aneurysms.
Aneurysms of the large arteries have been frequently reported as
having occurred in young children, even as early as in foetal life,
but in the present case not only was the child four years old, but the
aneurysms were several and their etiology obscure. Stress has been
laid by many on the effect of septic emboli in weakening the artery
wall by inflammation spreading from the infective clot in causing
H7
aneurysms in children suffering from septic endocarditis, but no such
cause was operative in this case. Syphilis is also commonly
considered a cause, but though no Wasserman blood test was made
on this boy, it is extremely improbable that syphilis was present.
Though there was a history of a fall antecedent to the appearance
of one of the tumours in this case the relationship of the two things
as cause and effect was not definite, while in the cases of the neck
tumours any competent injury must have been obvious at the time
it occurred. As causes, it appears that septic emboli, syphilis and
injury were not present in this case.
At one time, the presence of four arterial dilatations in a child of
four years old would have been held to justify the non-illuminating
diagnosis of 1 aneurysmal diathesis/
REFERENCE
Macfie, and Ingram (1920). Three cases of Cardiac Aneurysm in Native Boys of the Gold
Coast. Ann. Trop . Med. and Parasitol ., Vol. XIV, p. 147.
LAPPETED ANOPLOCEPHALA IN
HORSES
BY
WARRINGTON YORKE
AND
T. SOUTHWELL
(Received for publication 22 ]uly t 1921)
Plate XVII.
I. ANOPLOCEPHALA RHODESIENSIS, nom. nov.
About two hundred and fifty specimens of this parasite were
collected by one of us in 1912, from eight zebra (Equus burchelli )
in North Eastern Rhodesia.
TECHNIQUE. Most anatomical details were easily elucidated by
the following procedure : —
The worms were stained, en masse , for several days in dilute
acetic-acid-alum carmine, then washed and taken through the
alcohols into clove oil. As soon as the worm was clear, it was placed
under a binocular microscope and segments were detached, one at a
time, by means of a surgical needle or cataract knife, beginning with
the posterior segment. The segments were then mounted serially
with the anterior surface upwards. This procedure was quite simple
until about segments 10 to 15 were reached, when it was found that
the segments were so small and close together that the detachment of
single ones was difficult and tedious, and also often unsatisfactory.
Anterior to this point sections were therefore cut with a microtome,
when required. Horizontal sections were necessary, however, to
determine the structure of the uterus, e.g. } the presence, or absence,
of anterior and posterior outpocketings, and certain other details.
External Anatomy.
The largest specimen measured 114 mm. long, 22 mm. broad and
3*25 mm. thick, and the smallest 14 mm. long, 4 25 mm. broad and
250
r /5 mm - thick. Specimens less than 30 mm. long were immature.
Most of the larger worms approximated the following dimensions : —
Length 90 mm., breadth 20 mm., thickness 3 mm.; number of
segments, about two hundred and fifty.
The following measurements show the size of our largest worm
as compared with measurements made by^Fiihrmann and Collin of
the same species : —
Fuhrmann ...
Collin
Our specimen
Long Broad
... 50 mm. 18 mm.
... 70 mm. 26 mm.
... 114 mm. 22 mm.
Thick
3 mm.
5-6 mm. ?
3.25 mm.
The smallest specimen contained ninety segments, and the
posterior extremity was much narrower than the middle of the worm,
and was rounded; the last few segments were longer (0 25 mm.) than
the posterior segments in full-sized worms. A considerable number
of Oestrid larvae (Gastrofhilus sp.) were found firmly attached to
many of the worms.
Head . The head is always conspicuous; it is cuboid, with a
truncate anterior extremity; the breadth is considerably greater than
the length. In large worms measuring 90 mm. the head is about
2 mm. long and 3*25 mm. broad.
Suckers . The suckers, which are situated on the anterior surface
of the head, are directed straight forward. They are about 0*7 mm.
in diameter, and their cups are surrounded by a definite muscular
rim having a diameter of about 200 j*. The suckers were separated
from one another by more or less well marked grooves.
Lappets . Immediately behind each sucker is a large lappet.
The length and breadth of the lappets varies considerably, possibly
owing to contraction during fixation, but when they are fully
extended they measure about 1*25 mm. long, 1*75 mm. broad, and
the distance between the bases of the two dorsal or two ventral
lappets is about 0 6 mm. In some specimens the lappets were
globular and filled with liquid.
Segments . The segments are very shallow and are imbricated.
Posteriorly, the worm increases gradually in size, the greatest
breadth being usually about 1 cm. from the posterior extremity; it
must be remarked, however, that the general shape of the worm
varies considerably in different individuals, possibly owing to
different degrees of contraction during fixation (Plate XVII).
251
Internal Anatomy.
Muscular system. Both the longitudinal and transverse muscles
are strongly developed, and in transverse sections of mature segments
the thickness of the former is about 130/1, and that of the latter
about 60/1 (fig. 3). The dorso-ventral muscle bands are not so well
developed. A short but powerful muscle connects the internal
extremity of the cirrus pouch with the transverse muscles of the
ventral surface.
Nervous system. There are three longitudinal nerves on each
side, the median being far the most prominent. The other two,
dorsal and ventral, are slightly lateral to the median nerve and close
to the transverse muscular layer (figs. 1 and 3).
Excretory system. The water vascular system is enormously
developed. On each side there are two vessels, a very large ventral
vessel having a diameter of about 35 fi, and a much smaller dorsal
vessel which appears to be interrupted from time to time. The
ventral vessel is internal to the dorsal vessel which lies over the
median nerve trunk. The remarkable development of the water
vascular system is one of the most striking features in sections of the
worm (figs. 2 and 3).
Genitalia. At least the whole of the posterior half of the worm
is sterile, exhibiting no trace of genitalia, and, as in none of the
specimens examined was there any evidence that segments had been
shed, it is probable that the worm is passed entire in the faeces.
Testes. These first appear about segment 4 or 5, and they
disappear about segment 25. They attain their maximum develop¬
ment between segments 11 and 15, where each testis measures from
about 55/* to 90 fi by 30 /* to 70 fi. They occupy the entire medullary
parenchyma between the ventral excretory vessel on one side, and
the cirrus pouch on the other; they never cross the ventral excretory
vessel. They usually lie three or four deep in the dorso-ventral
direction, but the larger ones may be only one or two deep (fig. 1).
Vas deferens. After running laterally from the testes for a short
distance, the vas deferens dilates into the outer seminal vesicle,
which usually lies ventral to the cirrus pouch, although it was
observed occasionally to lie dorsal or median to the latter structure.
It then narrows and enters the cirrus pouch, where it again dilates
to form the inner seminal vesicle. The cirrus pouch is remarkably
252
long, and passes dorsal to the ventral water vessel and median nerve
to reach the edge of the segment (figs. 1 and 2). The cirrus is long,
slightly coiled, and is armed with very minute spines. Packets of
spermatozoa of various shapes and sizes, but usually oval with
pointed extremities, are abundant in both the outer and inner seminal
vesicles. In the majority of mature segments, the measurements of
these various structures are approximately as follows: —
Length of outer vesicula seminalis
. 55°j“
Greatest breadth of vesicula seminalis ...
. 15 °/*
Length of inner vesicula seminalis
. 70 Ofi
Greatest breadth of vesicula seminalis ...
. 200/4
Total length of cirrus pouch .
. 1 , 000 fl
Greatest breadth of cirrus pouch
. 100/4
DORSAL
v.
Fig. 1. A. rbodesiensis. Segment, viewed anteriorly, showing male genitalia. r.r., cirrus
sac ; e.v.s., external vesicula seminalis; i.v.r., internal vesicular seminalis; /.m., longitudinal muscles;
/.*., lateral nerves ; median nerve ; r.j., receptaculum seminis; /., testes ; v ., vagina ; v.e.z ?.,
ventral excretory vessel; v./., vitelline glands, x 40.
Ovary. This first appears about segment 24 and disappears
about segment 34; it attains its maximum development about
segment 30. The poral wing has a lateral diameter of about 1 mm.
and the aporal wing of 2*2 mm. The median axis of the ovary is
slightly cn the pore side. The ovary consists of a series of vertical,
club-shaped columns arising from a ventral horizontal base. The
largest column measures about 170 fi dorso-ventrally and 60 fi
laterally. The columns decrease in size towards the periphery of
the ovary (fig. 2). In the antero-posterior direction they ate never
more than two deep.
*53
DORSAL
VENTRAL
Fig. 2. A. rbodesiensis. Segment, viewed anteriorly, showing female genitalia, c., cirrus;
c.j., cirrus sac ; e.v.s. y external vesicula seminalis; l.m., longitudinal muscles; lateral nerves;
m.ft.y median nerve ; 0., ovary ; r.r., reccptaculum seminis; f.m., transverse muscles; v ., vagina ;
v.e.v.j ventral excretory vessel; v.g., vitelline glands. X 15.
Receptaculum and vagina . In segment 16 the vagina is well
defined. From the pore it runs as a narrow tube ventral to the
cirrus pouch for a short distance, then crosses it posteriorly to reach
the dorsal surface. At this point it dilates and runs inwards just
below the dorsal transverse muscle fibres, to enter the large
receptaculum seminis. The receptaculum seminis is roughly club-
shaped, with a greatly dilated internal extremity which is bent upon
Fig. 3. A. rbodesiensis. Aporal extremity of segment, viewed anteriorly, showing uterus,
longitudinal muscles, nerves and excretory vessels, /.m., longitudinal muscles; /.w., lateral nerves j
m.n.j median nerve ; /.m., transverse muscles; h., uterus ; r.e.r., ventral excretory vessel, x 40.
254
itself and almost reaches the ventral transverse muscle fibres. The
vagina and receptaculum are always loaded with packets of
spermatozoa (figs, i and 2). From its median surface there arises
a small tube—the fertilisation canal—which runs dorsally, receiving
the ducts of the shell gland, vitelline glands and ovary, and finally
opens into the uterus (fig. 4). The genital pores are all dextral.
Vitelline glands. Appear for the first time about segment 7
and are mature about segment 40. They lie ventrally, close to the
internal extremity of the receptaculum seminis (fig. 2), and consist
of two wings which do not appear to be lobular as in some species
of Anoplocephala .
Shell gland. As far as could be ascertained, it lies dorsal to the
vitelline glands and is almost always obscured by them.
Uterus. Visible for the first time about segment 15 as a cell¬
string running across the segment. In segments 40 to 70 or 80 it is
DORSAL
Fig. 4. A. rbodesiensis . Diagrammatic representation of fertilisation canal and connecting
ducts, f.c.j fertilisation canal; o.d oviduct; r.s. } receptaculum seminis; «., uterus; v.J.,
vitelline duct. X 250.
well developed and contains ova; at this point it is a straight wide
tube devoid of outgrowths, it does not occupy the whole of the
dorso-ventral diameter of the segment, nor does it extend laterally
beyond the ventral excretory vessel (fig. 3). Further development
beyond this stage was not seen, all the posterior segments being
sterile.
Eggs. Notwithstanding the fact that about sixty worms were
examined, no mature eggs were found. Eggs varying in size from
12 fi to 25 /jl were often found in the same uterus. The largest eggs
*55
seen had apparently three envelopes, the outer measuring about
25 n, the middle about 20 fi, and the inner, which was completely
filled with the embryo, about 11 /*. These were, however, very rare,
and by far the largest number of eggs seen measured about 12 fi to
15/* in diameter. They contained large oil and yolk globules. All
attempts to discover a pyriform body failed.
Scattered amongst the eggs in the uterus there occurred large
numbers of other cellular structures apparently of a nutritive
character (fig. 5). They varied in size, within wide limits, and
many seemed to be in process of degeneration.
Fig. 5. A . rbodesiemis . Eggs and nutritive cells, e., egg; n . c ., nutritive cells, x 730.
DIAGNOSIS. The following are the chief diagnostic characters : —
1. Presence of lappets.
2. Its large size and numerous segments.
3. The enormous development of the water vascular system
and longitudinal musculature.
4. The large number of sterile segments.
Abildgaard, in 1789, described a cestode from a horse, to which
he gave the name Taenia magna; this worm had no lappets behind
the head. Zeder, in 1800, also described an equine tapeworm,
which had no lappets behind the head, to which he gave the name
T. plicata . Rudolphi, in 1808, referring to T. zebrae y Sander,
collected from a zebra, placed it amongst the species dubiae , and
stated that it had affinities with T. plicata. Presumably, therefore,
256
T. zebrae of Sander did not possess lappets behind the head.
Later, Cobbold expressed the opinion that T. plicata was a synonym
of T. magna , Abildgaard.
The genus Anoplocephala was established by E. Blanchard
in 1848.
Collin (1891) gave a general description of a worm collected from
an African zebra which resembled A . perfoliata in possessing lappets
behind the head. He named the species Taenia zebrae .
Gough (1908) states that A. magna (Zeder), var. pediculata ,
Railliet, were found by him in the horse, donkey and zebra, and that
this species is not so rare in S. Africa as elsewhere. He points out
that as Collin’s worm possessed lappets, it must be distinct from
T. zebrae , Sander, and is, therefore, a species without a name.
Fiihrmann (1910) gave a careful description of Collin’s species
of T . zebrae from material obtained by the Kilimandjaro expedition,
but did not deal with the synonymy, nor did he refer to Gough’s
paper. Fiihrmann has been kind enough to send us a specimen of
his A. zebrae , which we have examined, and which is undoubtedly
the same species as the one described by us. It should be noted,
however, that Fiihrmann in his description fails to mention the
interesting fact that the greater part of the worm is sterile (at least
the posterior half), and furthermore, that he describes and figures
ripe eggs furnished with a pyriform apparatus. These have never
been seen by us, notwithstanding the fact that we have examined,
minutely, the uterus from the ripest segments of over twenty of our
specimens and also that of the specimen Fiihrmann was good enough
to send us.
Douthitt, in 1915, stated that A. plicata (Zeder) and A. zebrae
are synonyms of A. magna (Abildgaard), and draws no distinction
between Sander’s T . zebrae and Collin’s T . zebrae. He was
apparently unaware of Fiihrmann’s paper, as he makes no reference
to k.
It is clear that the worm which Collin named T. zebrae differs
from Sander’s T . zebrae , which is apparently identical with
T. magna , Abildgaard, in that Collin’s species is lappeted, whereas
Sander’s is not. T. zebrae 9 therefore, is a synonym of T . magna ,
and Collin’s worm is, as Gough says, a species without a name. We
propose to designate it A. rhodesiensis.
*57
II. ANOPLOCBPHALA PERFOUATA (Gocze, 1782),
Blanchard, 1848.
The museum of this School contains the following collection of
lappeted Anoflocephala from horses and mules: —
Horse A. Three specimens from Chesterfield, collected by A. W. Noel
Pillers, December, 1909.
Horse B. Eight specimens from Chesterfield, collected by A. W. Noel
Pillers, July, 1910.
Horse C. Four specimens from Sheffield, collected by A. W. Noel
Pillers. No date.
Horse D. Two specimens from Manititlan, Mexico. No further informa¬
tion.
Mule E. Ten specimens from Argentine, collected in 1917.
As a result of preliminary examination of the anatomy of
specimens from the above sources, we reached the conclusion that a
number of different species* were represented. On further examina¬
tion, however, we were impressed with the fact that great variations
occurred not only in worms from the same source, but even in
different segments of the same specimen. This led us to the
conclusion that the differences which at first we considered to be of
specific value were of inconstant occurrence, and consequently that
A. perjoliata (Goeze, 1782) is a species which exhibits considerable
variation.
In this paper we propose to re-describe the worm, drawing
attention to the variations which may occur.
External Anatomy.
The worms from the different sources were approximately the
same size, varying from about 30 mm. to 45 mm. in length, except
in the case of specimens from Horse B, which were fixed in a very
extended condition and were of a gelatinous consistency. These
were considerably longer, varying from 44 mm. to 70 mm. As a
rule, the worms attain a maximum breadth of about 12 mm.
Head . This is prominent and almost cubical in shape; the
length (2*5 mm.) is nearly equal to the breadth (3 mm.). In the
specimens from Horse A the head was, however, distinctly broader
(2*75 mm.) than long (1*5 mm.). The suckers and lappets resemble
closely those of A. rkodesiensis .
258
Segments. The number of segments in an adult worm varies
from about ninety to one hundred and thirty. The shape of the
worm varies enormously, as will be seen in the photographs. Except
that the worm is much less massive and consists of fewer segments
than A. rhodesicnsis , it exhibits no constant external difference from
the letter species (Plate XVII).
A M B del
Fig. 6 . A. perfoliata. Ventral view of anterior portion of a worm from Horse B ; cleared in
carbolic acid, c.s ., cirrus sac j /., lappets; median nerve; 0., ovary; r.r., receptaculum
seminis; testes ; uterus; v.t.v., ventral excretory vessel; 1vitelline glands. X 6.
Internal Anatomy.
Muscular system . As in A. rhodesiensis , except that the longi¬
tudinal fibres are not so strongly developed.
'Nervous system. The number of longitudinal nerves on each
side varied. As a rule, there were three on each side, but in
specimens from Horse B only the. median nerve was constantly
present, although the dorsal and ventral nerves were also seen in
some segments. This variation was observed in the different
259
segments of the same worm, with the result that some segments
exhibited only one nerve on each side, whilst in others three nerves
were found on both sides, and in still others three nerves on one side
and one on the other.
Excretory system . This differed from that of A. rhodesiensis
only in that the vessels were not so large.
Genitalia . The segments become increasingly ripe towards the
posterior extremity, and only a few sterile segments were found
scattered here and there. This is in striking contrast to the long
chain of sterile segments forming the posterior half of A. rhodesiensis .
Moreover, examination of the posterior extremities of worms
from various horses left no reason to doubt that segments are shed.
Fig. 7. A. perfoliata. Segment, viewed posteriorly, showing male genitalia. r.i., cirrus sac ;
e.v.s.y external vesicula seminalis; t.vj., internal vesicula seminalis; longitudinal muscles;
lateral nerves ; m.h., median nerve ; 0 ., ovary; r.r., receptaculum seminis ; testes ;
transverse muscles; v., vagina ; v.e.v., ventral excretory vessel, x 20.
Testes . These first appear about segment 12 and disappear
about segment 30. They attain their maximum development
between about segments 17 and 22. In all the specimens examined,
excepting those from Horse B, the fully developed testes occupied
the whole parenchyma between the aporal excretory vessel and the
cirrus pouch (fig. 7). In Horse B, whilst this state of things was
occasionally found, the testes, as a rule, were fewer in number and
more limited in distribution, being congregated mainly in the dorsal
i 6 o
portion of the parenchyma; they did not extend laterally nearly so
far as the aporal water vessel. There appears to be no doubt that in
old worms the testes degenerate and entirely disappear. No trace
of testes were found in the worths from Horse A.
Vas deferens . The appearance of the internal and external
seminal vesicles varied considerably, and they did not exhibit any
constant relationship one to the other. The outer seminal vesicle
was usually ventral to the inner seminal vesicle, but this was not
invariably the case; sometimes the former lay directly internal to the
latter, or even dorsal to it.
Ovary . This first appears about segment 25 and disappears
about segment 45. It attains its maximum development about
segment 37. The poral wing has a diameter of about half that of
the aporal wing. In structure it resembles that of A. rhodesiensis
DORSAL
I m trn
Fig. 8. A . pcrfoliata. Segment, viewed posteriorly, showing female genitalia, c.v.s., external
vericula seminalis; i.v.s internal vesicula seminalis; l.m longitudinal muscles ; lateral nerves ;
m.n., median nerve ; o., ovary; r.s ., receptaculum seminis; t.m transverse muscles; r., vagina ;
v.e.v.y ventral excretory vessel; v.g. y vitelline glands, x 20.
(fig. 8). As in the case of the testes, the ovaries had entirely
disappeared in worms from Horse A.
Receptaculum and vagina . The vagina appears early and is
well defined in about segment 8. These structures resemble in all
respects those of A . rhodesiensis .
Vitelline and shell glands . Similar to those of A . rhodesiensis.
Uterus. The uterus appears very early, about segment 12, as a
delicate cell-string running across the segment. It gradually
26 i
enlarges and attains its full development only in the last few
segments, where it is a wide tube completely filling the medullary
parenchyma, hut only occasionally crossing the ventral excretory
vessels. In the fully developed uterus there are numerous anterior
and posterior outpocketings, so that in anterior or posterior views of
whole segments the uterus appears to be composed of a number of
separate compartments (fig. 9). Sections, however, showed clearly
Fig. 9. A. perfoliata. Segment, viewed posteriorly, showing fully developed uterus. e. } egg ;
l.m., longitudinal muscles; /.m., transverse muscles; u. f uterus. X 40.
that the organ consisted of a central cavity with numerous anterior
and posterior bulges between the dorso-ventral muscle fibres (fig. io).
To a certain extent there are bulges on the dorsal and ventral surfaces
of the uterus, but they are not so prominent as those on the anterior
and posterior surfaces. The degree of outpocketing varied in
Fig. io. A. perfoliata. Horizontal section of segment showing fully developed uterus. X 40.
different worms, sometimes being well marked and in other cases
less obvious. Of the worms examined by us, the outpocketing was
most marked in those from Mule E, and least marked in those from
Horse B. Occasionally a well-developed uterus was seen which
contained few or no eggs and a mass of debris.
Egg s . Fully mature eggs were, found only in the last few
segments. The diameter of the outer envelope was about 8oyu, and
that of the embryo about 16/x, whilst the length of the horns of the
pyriform apparatus is about 18/*. Yolk granules about 7/1 in
262
diameter occurred plentifully (fig. 11). As the egg matures, the
middle envelope gradually shrinks, and its measurement is therefore
of no value. In quite ripe eggs the horns of the pyriform apparatus
A.M.B.deU
Fig. 11. A. perjoliata. Eggs showing pyriform apparatus. X 360.
are prolonged into very long, slender filaments which eventually
unite with one another.
DIAGNOSIS. Fuhrmann, as a result of a comparison between his
own observations on the lappeted Anoplocephala from the zebra
(called by him A. zebrae ), and Kahane’s description of A. perjoliata ,
gives the following points of difference between the two worms.
Cirrus pouch. In A. zebrae it extends beyond the longitudinal
nerve and ventral water vessel, whilst in A. perjoliata it scarcely
reaches the water vessel.
Vesicula seminalis . In A. zebrae , it is dorsal or ventral to the
cirrus pouch, and in A. perjoliata it is posterior.
Ovary . Much more strikingly asymetrical in A. zebrae than
in A. perjoliata.
We have carefully examined these points, and have reached the
conclusion that they have no specific value. In the specimens of
A. perjoliata examined by us, the cirrus pouch extended over the
longitudinal water vessel and nerve to the edge of the segment, just
as it does in the worm from the zebra. The relative position of the
outer seminal vesicle and the cirrus pouch is inconstant, and finally
263
the aporal wing of the ovary is about twice the size of the poral wing
in both worms.
In our experience, the only constant points of difference between
the lappetcd Anoplocephala of the zebra and horse, viz.,
A. rhodesiensis and A. perfoliata , are as follows: —
(1) A. rhodesiensis is much more massive than, and has almost
twice as many segments as, A. perfoliata.
(2) The posterior half of A. rhodesiensis is entirely sterile,
whereas in A. perfoliata the segments become increasingly
ripe up to the posterior extremity of the worm.
REFERENCES
Beddard, F. E. (1911). Contributions to the Anatomy and Systematic Arrangement of the
Cestoidea. I : On Some Mammalian Cestoidea. Proc. Zool. Soc., London.
- (1911). Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.
II : On Two New Genera of Cestodes from Mammals. Proc. Zool. Soc., London.
- (1912). Contributions to the Anatomy and Systematic Arrangement of the Cestoidea.
IV : On a species of Inermicapsifer from the Hyrax, and on the Genera Zscbokkeella,
Thysanotaema , and Hyracotaenia. Proc. Zool. Soc., London.
Bischopt, C. R. (1912). Cestoden aus Hyrax. Zool. Anz ., 39.
Blanchard, R. (1891). Notices helminthologiques (deuxi^me series). Mbn. Soc. Zool.,
France , 4.
Collin, A. (1891). Parasiten aus dem Darm des Zebra. Si/*, des Ges. Nat. Fr. zu Berlin.
Deiner, E. (1912). Anatomia der Anoplocephala latissima (nom. nov.). Arb. Zool. Inst.,
Wien., 19.
Do uthi tt, H. (1915). Studies on the cestode family Anoplocepbalidae. Illinois Biological
Monographs, University of Illinois.
Fuhrmann, O. (1902). Die Anoplocephaliden der Vogel. Centralbl.f. Bakt., I, Abt., Orig. 32.
- (191°). Vermes. Wissenscbaftlicbe Ergebnisse der Scbtoediscben Zool. Exp. nacb
dem Kiliamndjaro dem Meru, 1905-6. Band 3, Abt. 15-22, Stockholm.
Gough, H. L. (1908). Notes on Some South African Parasites. South African Association
for the Advancement of Science, Grahamstown.
Hall, M. C., and Hoskins, P. H. (1918). The occurrence of Tapeworms, Anoplocephala
spp., of the Horse in the United States. Cornell Veterinarian , October.
Kahane, Z. (1880). Anatomie von Taenia perfoliata, Goeze, als Beitrag zur Anatomia der
Cestoden. Zeitscbr.f. toiss. Zool., 34.
Leidy, J. (1855). Notices on some Tapeworms. Proc. Acad. Nat. Sc., Philad., 7.
Neveu-Lemaire, A. (1912). Parasitologie des Animaux Domestiques, Paris.
von Linstow, O. (1901). Helminthen von den Ufem des Nyassa-Sees—ein Beitrag zur
Helminthen-Fauna von Siid-Afrika. Jen. Zeitscbr.f. Naturto., 35.
MacCallum, G. A., and MacCallum, W. G. (1912). On the Structure of T. gigantea.
Zoolog. Jahrb. Syst., 32.
Neumann's Parasites (1905). Macqueen’s translation, 2nd edition, London.
Railliit, A. (1893). Traiti de Zoologie medtcale et agricole, 2nd ed., Paris.
-Henry, A., et Bauche, A. (1914). Sur les Helminthes de L’E 16 phant d’Asic.
Bull. Soc. Path. Exot., Tome VII, Nos. 1, 2, and 3.
Ransom, B. H. (1909). Taenoid Cestodes of North American Birds. Bull. U.S. Nat. Mus ., 69.
Shipley A. E. (1900). A Description of the Entozoa, collected by Dr. Willey during his
sojourn in the Western Pacific. {In Willey, Zool. Results, Part 5, Cambridge.)
Stiles, C. W. (1895). Notes on Parasites, No. 38. Preliminary Note to 4 A revision of the
adult leporine cestodes.’ Vet. Mag.
- (1896). A Revision of adult tapeworms of hares and rabbits. Proc. U.S. Nat. Mus.
- and Hassall, A. (1902-1912). Index-catalogue of Medical and Veterinary Zoology.
Author’s Index. Bur. An. Ind. Bull., 39.
- (1912). Index-catalogue of Medical and Veterinary Zoology. Subjects : Cestoda
and Cestodaria. Hyg. Lab. Bull., 85.
Zschokke, F. (1888). Recbercbes sur la structure anatomique et bistologique des Cestodes , Genive.
EXPLANATION OF PLATE XVII
Fig. i. Photograph of Anoplocephala rhodesiensis showing bots
attached. Natural size.
Fig. 2. Photograph of Anoplocephala perfoliata from Mule E,
Horse B and Horse C, and of Anoplocephala rhodesiensis
from a Zebra. (In order from left to right.) Natural
size.
Annals Ttop. \Ied, & ParasitolVol . W
PLATE XVII
Fig. i
Fig. 2
C. Titiling £? Co., /./</., /m/>.
265
THE FEEDING HABITS OF STEGOMTIA
CALOPUS, Meigen
BY
R. MONTGOMERY GORDON
AND
C. J. YOUNG
From the Laboratory of the Liverpool School of Tropical
Medicine , Manaos
(Received for publication 28 July, 1921)
It has been stated by Marchoux and Simond (1906) that
Stegomyia calopus , under normal conditions, that is, while at
liberty, ceases to bite man during the day after the first six or eight
days of its existence in the perfect state, but that after the eighth
day it is sometimes observed to bite towards 6 p.m. before nightfall.
They also state that yellow fever is not contracted during the day,
but only ‘ k la chute du jour ou pendant la nuit.’ ‘ Nous avons en
effet constate experimentalement qu’a la p&riode de sa vie ou il
possede le pouvoir infectant, le Stegomyia fasciata en liberte ne
cherche pas a piquer Thomme entre 7 heures du matin et 5 h. £ du
soir. La transmission est done nocturne/ This they held to
account for the fact that the inhabitants of Petropolis when visiting
Rio only during the day did not contract yellow fever. In their
experiments, however, the mosquitoes were confined in a screened
room. Seidelin and Connal (1914) and Macfie (1915-16) showed
that S. calopus would bite at any hour irrespective of age, when in
captivity.
According to Marchoux and Simond (1906) the minimum
interval between the date on which S. calopus acquires the infection
and that on which it becomes infective is twelve days.
The following experiments were undertaken with a view to
investigating the feeding habits under as natural conditions as
possible. They were carried out in Mangos.
All the mosquitoes were hatched in the laboratory and kept in
wire gauze cages, males being always present. Each female was
removed from the stock cage in a glass tube and allowed to feed to
repletion on one of us during the hours of daylight. After feeding
they were 1 marked ’ by amputating the hindlegs through the tibiae,
266
but in a few instances some part of the femur was accidentally
removed. This method of ‘marking 1 is considered sufficiently
distinctive as wild specimens of Stegomyia , observed by us at other
times, have never shown this mutilation. After feeding they were
kept for not less than fourteen days, sugar being available as food,
and during the last two or three days of this period were again
given an opportunity to bite, also during daylight. They were
then released under the conditions described below and during the
four succeeding days, in the place where they had been released,
the mosquitoes biting us were observed for one hour each during the
day and night. The observations at night were made by electric
light. No unmarked females were released along with the 1 marked.’
Three experiments were carried out.
Experiment /. Date: 23rd March to 16th April, 1921. Sixty
female Stegomyia were fed a first time and * marked ’ between
23rd and 28th March. Of these thirty-three survived, and were
offered a second feed on the nth and 12th of April; thirty fed and
three refused to feed. Two escaped after feeding. The remaining
thirty-one females along with twelve males were released at 7 p.m.
on the 14th April in a first-floor bedroom of about 14 ft. by 12 ft.
by 10 ft. This room was in use, the occupant using a mosquito net
at night; it had two unscreened windows which were constantly
open, and the mosquitoes also had free access to other parts of the
house. A glass jar containing water was placed in the room, but
no eggs were deposited in it up to the 23rd April. ‘ Marked ’
mosquitoes were released after feeding.
Observations were made as follows: —
Date .
1
April ...| 1
5
!
1
6
1
7
1
8
Time.
1 12.30
P.M. ...j to
1 «- 3 °
8.30
t°
9.30
12.30
to
1.30
8.30
to
9 - 3 °
12.30
to
1.3c
8.30
to
9 - 3 °
12.30
to
1.30
8.30
to
9 - 3 °
S. calopus I
biting ... ■!
* Marked ’... 3
. . |
0
1 2
1 •
0
0
0
0
0
t
Unmarked.. J 3
1
0
1 °
0
0
0
0
0
Note.— In addition, one of us was bitten by a ‘marked’
mosquito at 7.30 a.m. on the 15th.
267
Experiment //. Date: 5th to 29th April, 1921. Sixty
female Stegomyia were fed a first time and ‘ marked 9 between the
5th and nth April. Of these twenty-three survived. They were
offered a second feed on the 23rd and 24th April; twenty-one fed
and two refused. They were released with about twelve males at
5 p.m. on the 25th in a first-floor bedroom in a different part of the
town from Experiment I. This room was about 20 ft. by 14 ft. by
12ft. in size, was in use, the occupant using a mosquito net at
night, had four large windows constantly open, and the mosquitoes
had free access to other parts of the house.
Observations were made as follows: —
Date.
April
2 5
26
1
27 J
28
2 9
Time.
P.M.
O'
1
00
12 —I
1
12—I j
8-9
_ .
12—I
8-9
12—1
'
S. calopus
biting ... •
‘ Marked *...
0
0
O
0
O
0
O
0
Unmarked...
0
O
2
O
O
O
i_
0
0
Experiment III. Date: 1st May to 3rd June, 1921. Seventy
female Stegomyia were fed a first time and 1 marked * between
1st and 14th May. Of these fifty survived. Offered second feed
on 28th, 29th and 30th May; forty-three fed and seven refused.
These fifty were released along with about twelve males at 3 p.m.
on the 30th May in a ground-floor room behind the laboratory,
about 17 ft. by 10 ft. by 15 ft. in size and opening directly to the
outside. This room had one window, the upper part of which was
imperfectly screened and the lower part had wooden slats, the
apertures between these allowing free passage to mosquitoes. The
door was open all day and for one hour at night when observations
were made. Otherwise the room was not used at night. A jar
of water was placed in the room, and some eggs were found on
31st May. ‘Marked* mosquitoes were released after feeding.
268
Observations were made as follows: —
Date .
May
3 o
3 *
June 1
2
3
1—2
Time.
P.M.
8.30
to
9.30
1—2
8.30
to
9 - 3 °
1—2
8.30
to
9 - 3 °
1—2
8.30
to
9.30
5 . calopus biting
‘ Marked ’...
!
3
7
3
*
0
1
0
0
Unmarked...;
1
1
4
4
1
3
0
0
5
Note. —Each of us was bitten in the laboratory by a ‘marked*
Stegomyia on the 7th June, one at 9 a.m. dnd the other at 4 p.m.
SUMMARY
Experiment I. Thirty-one ‘ marked * female Stegomyia were
released at night not less than fourteen days after their first blood
meal. During the succeeding four days, five ‘ marked * and three
unmarked fed during daylight. None were observed to feed at
night.
Experiment II. With the exception of two unmarked Stegomyia,
none were observed to feed.
Experiment III. Fifty ‘marked* female Stegomyia were
released during daylight not less than fourteen days after their first
blood meal. During the succeeding four days, ten ‘ marked * and
ten unmarked fed during daylight, and six ‘marked* and eight
unmarked fed during the night.
CONCLUSION
Stegomyia calopus females will bite either by day or night, over
fourteen days after their first blood meal, while under no artificial
restraint and having opportunities of selecting day or night for
feeding.
REFERENCES
Macfii, J. w. s. (1915-16). Bulletin of Entomological Research , Vol. VI.
Maichoux and Simond (1906). Annales de Vlnstitut Pasteur Vol. XX.
SiiDELiN and Connal (1914). Telloto Fever Bulletin , Vol. Ill, No. 3.
269
NOTE ON A CASE OF LEPROSY
BY
WARRINGTON YORKE
AND
S. ADLER
(Received for publication 2 August , 1921)
PLATE XVIII
Native of Hong Kong, age 45. Came to the outpatient
department of Tropical School, 29th July, 1921, complaining of
feeling ill, ulceration of face and generalised skin eruptions.
Present Condition. The appearance of the face at once
suggested leprosy. Both alae nasi were swollen and ulcerated,
and there were fairly well developed nodules on the chin, forehead
and cheeks. There was an extensive raised, ham coloured, rash on
the arms, forearms, trunk and legs. The affected areas of skin were
not anaesthetic except for small patches on.the inner side of the left
leg and foot and on the outer side of the right leg. No thickening
of the superficial nerve trunks was observed. Scrapings from the
skin lesions and from the alae nasi showed very numerous lepra
bacilli .
HISTORY. The most interesting feature of this case is its history.
The patient arrived in England in 1912 in good health, and has*
been employed until a short time ago as a laundryman in Cardiff.
It was not until 1916—four years after coming to England—that he
first noticed small spots on the left side of the face. No further
change was observed until twelve months ago, when the ham-
coloured eruption appeared on the trunk and upper and lower
limbs. Five months ago he noticed for the first time the swelling
of the alae nasi .
There is no evidence that the case had been diagnosed before we
saw him: the patient and his friends were quite unaware of the fact
that it was leprosy.
270
EXPLANATION OF PLATE XVIII
Photograph of a case of leprosy showing swelling and ulceration
of alae nasi and nodules on various parts of face.
Annals Trop. Med. & ParasitolVol. XV
PLATE XVIIl
C. Titiling & Co., Ltd., Imp.
271
NOTES ON SOME FUNGAL INFECTIONS
IN WEST AFRICA
BY
J. W. S. MACFIE
(Received for publication 24 August , 1921)
CONTENTS
PAGE
I. A Fungus or the Genus Monilia Recovered from a Patient Sukfering from
Dysenteric Diarrhoea. 272
II. Two Fungi or the Genus Monilia Recovered from Africans Suffering from
a Peculiar Form or Diarrhoea . 275
III. A Note on a Case or Otomycosis. 279
IV. Tonsillar Nocardiomycosis ... ... ... ... ... ... ... ... 282
Fungal infections are abundant in West Africa, but hitherto
comparatively little attention has been devoted to them. Brief
notes are given in this paper on a few cases which have recently
come under our notice at Accra in the Gold Coast.
The part played by the fungi of the Genus Monilta in the diseases
from which the patients were suffering is obscure. Such fungi are
undoubtedly abundant in the tropics, and contamination with
them must be guarded against; it is also true that they are not
infrequently present in small numbers in the faeces of apparently
healthy individuals. Their occurrence in great quantities is,
however, a different matter. No evidence is brought forward in
support of the view that the species of Monilia described in this
paper were the sole cause of the dysentery and diarrhoea with which
they were associated, indeed there was evidence that the cases
referred to in Section II were due primarily to a dietetic deficiency,
but it is maintained that they were the cause of some of the
symptoms. For this reason, and because the intestinal disturbances
which permit of their multiplication may be of profound significance,
their recognition and investigation is of importance.
2J2
I. A FUNGUS OF THE GENUS MONIUA RECOVERED FROM
A PATIENT SUFFERING FROM DYSENTERIC DIARRHOEA
Towards the end of the year 1920 there occurred at Accra a
considerable number of cases of dysenteric diarrhoea, the cause of
which was obscure. From one such case, a European man about
forty years of age, the Monilia was isolated, of which the following
is a short description. The sample of faeces from which the fungus
was recovered contained much blood and mucous, but no amoebae
and no other parasite to which a pathogenic rdle could be assigned;
it contained, however, a considerable number of yeast-like cells.
From the same case, a week later, Dr. J. F. Corson isolated a
bacillus which gave the reactions of Morgan’s bacillus.
'Several species of the Genus Monilia have been isolated from
human faeces in various parts of the world, but especially in the
tropics, and some of them have been regarded as being the cause,
either directly or indirectly, of intestinal disease. Similar fungi
are frequently found in small numbers in specimens of faeces in West
Africa, and in drarrhoeic stools are sometimes very numerous.
With regard to the species isolated from this case of dysenteric
diarrhoea at Accra it is not possible to say if it was pathogenic, but
it may be noted that it closely resembled M. trofncalis , one of the
species which has been found to be present in sprue (Castellani,
1920).
The Organism. The Monilia found in this case was isolated
in a culture of the faeces on neutral-red lactose bile-salt agar. It
was Gram-positive and not acid fast. It grew well on most solid
media, and on glucose agar produced within twenty-four hours
an abundant creamy-white growth. Under anaerobic conditions
growth was less abundant and less rapid. It grew freely both at
37 0 C. and at the temperature of the laboratory (about 26° C.).
Gelatin and blood serum were neither liquefied nor stained. In
broth and peptone water a whitish deposit was thrown down,
whilst the media themselves remained clear; in both a surface pellicle
was formed. It produced an abundant white growth on potato,
which later developed a white efflorescence; the medium was not
stained. On solid media the growth was mainly composed of yeast¬
like cells, but a few branched septate hyphae were usually present;
in fluid media the hyphae sometimes predominated.
273
Its qualitative bio-chemical
reactions may be
tabulated as
follows, the symbols
representing:—A = acidity
; G = gas;
s = slight; O = neither acid nor
gas.
Arabinose
0
Inulin .
0
Rhamnose (isodulcite)
0
Amygdalin
0
Galactose
... AGs
Helipn.
0
Glucose.
... AG
Phlorrhizin
0
Laevulose
AG
Salidn .
O
Mannose.
0
Glycerol
O
Lactose ...
0
Erythrol
0
Maltose.
... AG
Adonitol
O
Saccharose
... AGs
Dulcitol.
0
Raffinose.
O
Inosite.
0
Amylum ...
... , O
Mannitol
O'
Dextrin.
O
Sorbitol.
0
Glycogen
O
If the cultures were kept, the acidity produced in the five sugary
media indicated tended to be superceded by alkalinity; this change
began to occur early, usually in less than a week. No change was
produced in Litmus milk at first, but after a week or ten days a
slight alkalinity was sometimes noticed; no clot was formed, and
the medium was neither decolourized nor peptonized. Indol was
not produced in peptone water.
As gas was produced in Glucose, Laevulose, Maltose, Galactose,
and Saccharose the organism comes into the fifth group of species
of Monilia , the Tropicalis group, according to the classification of
Castellani and Chalmers (1919). The more important bio-chemical.
reactions of the species in this group are shown in Table I. It will
be noted that the species here described differs in its reactions in
Mannitol and Dextrin from M. enterica , M. insolita > M. para -
tropicalis , and M . pulmonalis. From M. faecalis it may be
distinguished by the fact that it does not decolourize Litmus milk,
and does not stain blood serum brown; and from M . metatropicalis
by the fact that it does not clot milk. As regards M. nivea , the
fact that neither acid nor gas is produced in Raffinose is a point of
distinction, and it may be noted also that our species produces only
a slight amount of gas in Galactose and forms a pellicle on the
surface of broth, reactions which are not found in the case of
M. nivea . Our organism closely resembles M . tropicalis , the
greatest divergence being *in the reaction in milk. It is said that
by M. tropicalis ‘Litmus milk is generally rendered acid but is not
274
clotted,’ a statement which would seem to imply that the production
of acid is not constant nor characteristic. If this is so, the two
species appear to be identical, excepting that in the case of our
organism a pellicle is formed in broth.
Table I.
Species of the Genus Monilia belonging to the Tropicalis group.
Specie* of Monilia
Litmus milk
Glucose
Laevulose
Maltose
Galactose
Saccharose
Mannitol
Dextrin
Raffinose
§
c
3
(9
<
Broth
M. enterica
O/Alk
AG
AG
AG
AG
AG
As
As
O
O
c
M.faecalis .
A/DPs
AG
AG
AG
AGs
AGs
O
O
O
O
c
M. insolita .
As/Alk
AG
AG
AG
AG
AG
As
O
O
O
c
M. metatropicalis .
AC
AG
AG
AG
AG
AG
O
O
O
0 .
c
M. nivea .
O/Alk
AG
AG
AG
AG
AGs
O
O
AG
0 1
c
M. paratropicalis .
As/Alk
AG
AG
AG
AG
AG
O
Avs
O
0
CTP
M. ptdmonalis .
O/AlkD
AG
AG
AG
AGs
AG
Avs
O
A
AGs
CTP
M. tropicalis .
A
AG
AG
AG
AGs
AGs
O
O
O
O
C
Accra case : sputum and pleural
cavity ( M. accraensis )
O/Alk
AG
AGs
AGs
AG
AG
O
O
O
O
C
Accra case : dysenteric faeces ...
O/Alk
AG
AG
AG
AGs
AGs
O
O
O
O
CTP
A = acidity; Aik = alkalinity; C = clot (milk), clear (broth) ; CTP = clear, then pellicle ; D = decolourized;
G = gas; O = neither add nor gas ; P = peptonized ; • = slight; vs = very slight.
Finally it may be noted that this organism closely resembles a
species of Monilia {M, accraensis) recently found by us in the sputum
and pleural cavity of a patient suffering from tuberculosis at Accra
(1921). The chief points of distinction are that the intestinal species
produces only a slight amount of gas in Galactose and Saccharose,
and forms a pellicle on broth; whereas the pulmonary species
produces much gas in Galactose and Saccharose but only little in
Laevulose and Maltose, and does not form a pellicle on broth.
SUMMARY
From the faeces of a European with dysenteric symptoms a
fungus of the Genus Monilia was isolated. This organism belongs
275
to the Tropicalis group, and appears to resemble most closely
M. tropicalis (Castellani, 1909).
REFERENCES
Castellani, A. (1920). Milroy Lectures. Joum, Trop. Med. and Hyg., XXIII, p. 120.
- and Chalmers, A. J. (1919). Manual of Tropical Medicine. Bailliere, Tindall and
Cox, London, pp. 1082-1083, and p. 1086.
Macpie, J. W. S., and Ingram, A. (1921). Bronchomoniliasis Complicating Pulmonary
Tuberculosis in a Native of the Gold Coast, West Africa. Annals Trop. Med. &
Parasitol ., XV, pp. 53-58.
II. TWO FUNGI OF THE GENUS MONiLiA RECOVERED FROM
AFRICANS SUFFERING FROM A PECULIAR FORM OF DIARRHOEA
There occurs among the natives at Accra a well defined form
of persistent diarrhoea which has been considered by some of the
medical practitioners in the Gold Coast to be akin to sprue. The
stools are bulky, frothy, and generally canary-yellow coloured; the
tongue is red, fissured, and eroded; and the patients are wasted—
a condition which is often masked by oedemas of the limbs and by
ascites. The faeces in such cases usually teem with yeast-like cells
which on cultivation are found to be stages of fungi of the Genus
Monilia. The following are'short descriptions of the organisms
isolated from two such cases.
First Case.
The patient, a native man about twenty-seven years of age, was
suffering from an obstinate diarrhoea, and, was weak and wasted.
His tongue was red, irregularly fissured, indented by the teeth, and
partially eroded; the throat also was red. The faeces were canary-
fellow coloured and frothy; yeast-like cells were extremely abundant
in them, and Entamoeba colt and Blastocystis enterocola were also
present.
The Organism . The Monilia in this case was isolated from a
culture of the faeces on neutral-red lactose bile-salt agar. It was
Gram-positive and not acid fast. It grew well on most solid media,
and on Glucose agar produced within twenty-four hours an abundant
white growth, very fluid in consistence, and with a dull pellicle-like
surface. The yeast-like cells, which composed the greater part
of the growth, were oval and somewhat elongated; the average
2 j6
measurements of ten cells were, length 4*5/1, breadth r6 fi. Under
anaerobic conditions growth was less abundant and less rapid.
Gelatin and blood serum were neither liquefied nor stained. In
broth and peptone water a whitish deposit was thrown down, and
the medium remained clear; in broth a surface pellicle was formed.
On potato a dull white growth was developed which later showed a
white efflorescence; the medium was not stained. On solid media
the growth was mainly composed of yeast-like cells, but a few
branched, septate hvphae were also present; in fluid media hyphae
sometimes predominated.
The qualitative bio-chemical reactions of this Monilia may be
tabulated as follows: —
Arabinose
0
Inulin ...
. 0
Rhamnose (isodulcite) ...
0
Amygdalin
. 0
Galactose
0
Helicin ...
. As
Glucose ...
AG
Phlorrhizin
. O
Laevulose
AG
Salicin ...
O
Mannose...
O
Glycerol
. O
Lactose ...
O
Erythrol
. 0
Maltose ...
AGs
Adonitol
. O
Saccharose
AG
Dulcitol...
. O
Raffinose...
AGs
Inosite ...
. O
Amylum
O
Mannitol
. O
Dextrin ...
AG
Sorbitol ...
. O
Glycogen
O
The symbols representing : A = acidity ; G = gas; s = slight;
O = neither acid nor gas.
In the media in which acidity was produced the reaction tended
after a few days to be succeeded by alkalinity. No change was
produced in Litmus milk- Indol was not produced in peptone
water.
As gas was produced in Dextrin as well as in other media the
organism comes into the ninth group of species of Monilia , the
Pseudolondinensis group, according to the classification of Castellani
and Chalmers (1919). Two species belong to this group, namely,
M. pseudolondinensis , Cast., and M. pseudolondinoides , Cast.,
which appear to differ only in their actions on Litmus milk. The
more important bio-chemical reactions of these species and of the
organism just described are given in Table II; it will be noted that
277
the latter species differs from the other two in its actions on
Galactose, Saccharose, and Raffinose. These differences are
important, and since the classification of these fungi* is at present
Table II.
Species of the Genus Monilia belonging to the Pscudolondinensis group.
Species of Monilia
Litmus milk
Glucose
Laevulose
Maltose
Galactose
Saccharose
Raffinose
Dextrin
M. pscudolondinensis, Cast.
O
AG
AG
AG
AG
O
o
AG
M. pseudolondinoidts. Cast.
AC
AG
AG
AG
AG
O
O :
AG
M. africana, sp.n. .
O
AG
AG
AGs
O
AG
i AGs 1
1
AG
A — acid ; G = gas; s — slight; O = neither acid nor gas; C = clot.
based on such bio-cheniical reactions, the organism described here
must be regarded as a new species. For it, therefore, the name
Monilia africana is proposed.
Second Case.
The patient, an adult native man, a Moshi, was suffering from
persistent diarrhoea accompanied by ascites and oedema of the face,
especially the left parotid area, the lumbar region, and the penis.
There was no oedema of the legs. The faeces were semi-fluid, pale
canary-yellow coloured, and frothy; they contained very numerous
yeast-like cells.
The Organism. The Monilia in this case was isolated from a
culture of the faeces on neutral-red lactose bile-salt agar. The
colonies of yeast-like cells were rather slow in appealing, being
detected first after forty-eight hours’ incubation. It was Gram¬
positive and not acid fast. It grew well on most solid media,
spreading as a thin film, and on Glucose agar produced within
twenty-four hours an abundant white growth. The yeast-like cells
which composed the greater part of this growth were, as in the
previous case, somewhat elongated. Under anaerobic conditions
growth was slower. Gelatin and blood serum were neither liquefied
nor stained; the growth on gelatin was slow. In broth and peptone
water a whitish deposit was thrown down, the medium remained
278
clear, and no pellicle was formed. On potato an abundant whitish
growth was produced, and the medium was not stained. On solid
media the growth was composed mainly of yeast-like cells, but a few
branched, septate hyphae were also present; in fluid media hyphae
sometimes predominated.
The qualitative bio-chemical reactions of this Monilia may be
tabulated as follows: —
Arabinose
• • • . . .
0
Inulin .
0
Rhamnose (isodulcite) ...
0
Amygdalin
0
Galactose
A
Helian.
As
Glucose ...
AGs
Phlorrhizin
O
Laevulose
AGs
Salidn .
As
Mannose...
0
Glycerol
O
Lactose ...
0
Erythrol
O
Maltose ...
A
Adonitol
O
Saccharose
0
Duldtol.
O
Raffinose...
O
Inosite.
O
Amylum
O
Mannitol
O
Dextrin ...
... ...
O
Sorbitol '.
0
Glycogen
...
O
The symbols representing : A = acidity ; G = gas; s = slight;
O = neither add nor gas.
In the media in which acidity was produced the reaction tended
after a few days to be succeeded by alkalinity. No change was
produced in Litmus milk. Indol was not produced in peptone
water.
As gas was produced in Glucose and Laevulose only, the
organism comes into the second group of species of Monilia , the
Krusei group, according to the classification of Castellani and
Chalmers (1919). Two species are included in this group, namely,
M. krusei , Cast., and M. parakrusei , Cast., which appear to differ
only in their actions on Litmus milk. The more important
bio-chemical reactions of these species and of the organism just
described are given in Table III; it will be noted that the latter
species differs from the other two in its actions on Maltose and
Galactose besides in several minor points. These differences,
according to the system of classification at present in vogue, are
sufficient to justify the erection of a new species, and, therefore,
although the number of species of the Genus Monilia is already
embarrassingly large, we propose for this organism the name Monilia
enterocola .
279
Table III.
Species of the Genus Monilia belonging to the Krusei group.
Species of Monilia
Litmus milk
Glucose
Laevulose
Maltose
Galactose
Saccharose
M. krusei, Cast.
O
AG
AG
O
O
O
M. parahrusei, Cast.
AC
AG
AG
O
O
O
M. enterocola , sp.n.
O
AGvs
AGs
A
A
O
A = acid ; G = gas; s = slight; ys = very slight j O = neither acid nor gas; C = clot.
SUMMARY
From the faeces of two Africans suffering from a peculiar form
of diarrhoea two fungi of the Genus Monilia were isolated, both of
which appear to be hitherto undescribed species. The one belongs
to the Pseudolondinensis group, the other to the Krusei group of
Castellani and Chalmers. The names Monilia africana and Monilia
enterocola , respectively, are proposed for these organisms.
III. A NOTE ON A CASE OF OTOMYCOSIS
Otomycosis is said to be not uncommon in the tropics. It may
be caused by a considerable number of different fungi, and according
to Castellani and Chalmers (1919), 1 if they grow superficially, they
cause no symptoms; but if they penetrate into the mucous membrane,
they give rise to itching, and sometimes to pain/
The patient whose case is the subject of this note was a European
lady who consulted Dr. C. V. Le Fanu at Accra in the Gold Coast
Colony, West Africa, on account of pain and. irritation in the ear.
The symptoms had existed for at least a month, but although they
caused no little discomfort, there was no deafness. On examination
Dr. Le Fanu found that the external auditory meatus was unusually
narrow, but widened out at its inner end so as to form a small
chamber immediately external to the drum of the ear. From a patch
on the vault of this chamber, close to the drum of the ear, the fungus
28 o
was growing and hanging down like a veil. It may be said at
once that the condition was rapidly relieved, and apparently cured,
by an application containing as its principal constituent salicylic
acid.
The fungus, w r hich grew readily on Sabouraud’s maltose agar
and was easily obtained in pure culture on this medium, appeared
to belong to the Genus Sterigmatocystis , Cramer 1859. Two
species of this genus have been found in cases of otomycosis, namely,
5. antacustica and S. nidulans , but the characters of both, as given
in the descriptions to which I have access, differ slightly from those
of the fungus isolated at Accra. The exact determination of the
species of such a fungus is a matter for a mycologist. I shall,
therefore, restrict myself to recording certain characters which may
enable those who have made a special study of these fungi to place
it more exactly.
Cultures. On Sabouraud’s maltose s\gar the growth of the fungus
was rapid and spreading; at first yellowish-white and felt-like, then
slightly fluffy with the development of upstanding conidiophores,
then speckled with dark brown points when, the conidiophores began
to develop their spores, and finally dark brown or almost black all
over, resembling a mass of soot. On Glucose agar the initial growth
at a temperature of 28° C. was almost white, with a somewhat
puckered and wrinkled surface, but within twenty-four hours became
yellowish, and began to show upstanding, white conidiophores. On
the following day the whole surface of the medium was covered by
a yellow, wrinkled, felt-like growth, on which were very many
conidiophores, some already dark brown in colour. On the third
day the whole surface of the medium looked as if it had been thickly
dusted with soot. Growth, therefore, was rapid and abundant at
28° C., the temperature of the laboratory; it was, however, even more
rapid at 37 0 C. In peptone water the fungus grew mainly on the
surface, but detached fragments in the fluid produced delicate
networks of hyphae which sometimes resembled puff-balls. Dark
spores were formed on the conidiophores at the surface. In glucose
peptone water acid was produced, but no gas. On blood serum
growth was rather slow, the colonies were white, and the black spore-
bearing conidiophores did not develop for a long time and were
relatively scanty. Eventually the medium was liijuefied. In a
gelatin stab culture there was no deep growth, and no liquefaction.
A white surface growth developed with dark sooty grains, and
later, just below the surface, there formed compact cerebriform
whitish masses of the fungus. On potato growth was rapid and
abundant; at first a yellowish felt-work, finally a sooty mass. In
Litmus milk acid and clot were produced. The growth w f as mainly
at the surface, was yellowish, and developed the usual dark brown
sooty appearance.
Mycology . All the growths showed septate, branched hyphae
of varying diameter. The hyphae appeared colourless when seen
with a microscope. The lengths of the interspaces were very
variable. In fluid media, such as glucose peptone water,
chlamydospores were numerous in the surface felt-like growth. The
conidiophores were erect. In cultures they were raised about
i mm. above the level of the medium, but in the ear of the patient
they appeared to be considerably longer. They were white on first
appearance, but darkened later. The diameter of the stem was
variable, but averaged about 14 fi. The head was almost spherical,
slightly broader than long, its diameters averaging 40 /jl and 37 fi
respectively; it was covered almost completely by sterigmata, only a
very narrow zone at the proximal end, at the insertion of the stalk,
being free from them. Both primary and secondary sterigmata
were present; the primary sterigmata were about 12 n long (average
of ten), the secondary about long (average of ten). There were
usually four secondary sterigmata on each primary sterigma. The
spores were dark brown in colour, and spherical in shape; diameter
41* to 5’2/u, average 4 5/ti.
Animal inoculations . Two wild Mus rallus were given intra-
peritoneal inoculations of an emulsion of a small fragment of the
original culture in normal saline. No ill-effects were observed to
follow. A little of the same culture was applied to the external
auditory meatus of a pouched rat ( Cricetomys gambianus ), a sheep,
and a small monkey ( Cercopithecus patas) after scarification, but no
otomycosis was produced.
REFERENCE
Castellani, A., and Chalmers. A. J. (1919). Manual of Tropical Medicine, Third Edition,
p. 2012. London : Bailliere, Tindall and Cox.
IV. TONSILLAR NOCARDIOMYCOSIS
One case of tonsillar nocardiomycosis has been met with at Accra.
The patient, a European man, about thirty-two years of age, showed
a number of small white concretions in crypts of both tonsils. The
throat was not inflamed, and there was neither pain nor discomfort;
but the presence of the concretions had been detected by the patient
at least a month previously, and he was worried about them as they
did not show any signs of disappearing.
Scrapings from one of the white patches consisted mainly of
fungal hyphae in short lengths. They were about i/i in diameter,
branched, and either Gram-positive or composed of Gram-positive
coccoid bodies and rods connected together by Gram-negative
strands.
Inoculations made on agar and glucose agar were unsuccessful.
Further particulars with regard to the fungus cannot, therefore, be
given.
This condition is referred to by Castellani and Chalmers (1919)
in their Manual of Tropical Medicine . The above case is recorded
here merely to draw attention to the fact that it occurs in West
Africa.
REFERENCE
Castellani, A m and Chalmers, A. J. (1919)- Manual of Tropical Med'nine y Third Edition,
pp. 1747-8. London : Baillicrc, Tindall and Cox.
283
A FUNGUS OF THE GENUS NOCARDIA
CULTIVATED FROM HEART BLOOD
BY
J. W. S. MACFIE
AND
A. INGRAM
(Received for publication 24 August, 1921)
The fungus here briefly described was cultivated from blood
withdrawn from the heart at the autopsy on a patient who died in
May, 1920, at Accra, of an obscure complaint. The notes of the
case are as follows: —
The deceased was a native man, about twenty-five years of age,
regarded clinically as possibly suffering from encephalitis lethargica,
and said to have been* ill six days before death. He was
stated to have shown paretic symptoms of the arms, and to have
had convulsive seizures during which there was opisthotonus.
Consciousness was lost temporarily during the illness, but was
recovered before death.
At the autopsy, which was made on the 14th of May, 1920, eight
hours after death, the body was found to be well nourished; post¬
mortem rigidity was present in the legs and arms but absent from
the neck. There was an old scar over the external aspect of the
right upper arm immediately above the elbow joint, an old circular
cicatrix about the size of a sixpenny piece over the left malar bone,
and a small punctured wound at the back of the neck on a line with
the thyroid cartilage: the latter wound was oozing blood, and may
have been inflicted after death when placing the body in the coffin.
On examining the internal organs, both lungs were found to be
affected with broncho-pneumonia: the left lung was adherent to the
284
parietal wall by recent fibrinous formations. Brain: meninges
somewhat congested; blood vessels in sulci engorged; no pus on
the surface nor at the base. Heart: blood fluid; no abnormality
noted. Cultures were made with blood obtained from the heart by
puncturing the organ after searing its surface and taking every
precaution to avoid contamination: in these cultures the Nocardia
was grown. Digestive track, spleen, kidney, and liver: congested,
but otherwise showed no apparent pathological condition.
The organism cultivated from the heart blood of this case grew
well on blood agar and ‘ nasgar * at 37 0 C. and at the temperature
of the laboratory, about 26° C., producing a somewhat slowly
spreading growth which was very firmly adherent to the medium.
The colonies were at first smooth and dome-shaped, but later
became puckered and opaque in the middle and radially striated
and semi-transparent at the periphery. After a few days an
efflorescence appeared on the older parts of the growth which in
the earliest cultures was abundant and blue, and in later sub¬
cultures was scanty and grey or white. The medium sublying
the growth was not stained. The organism grew both aerobically
and anaerobically, but slowly and rather feebly under the latter
conditions. The cultures had no distinctive odour.
When examined microscopically the growth was seen to be
composed of freely branching non-septatte hyphae about ifi or less
in diameter. In old cultures many of the hyphae were more or less
fragmented, and the ends of some of the filaments were slightly
thickened and club-shaped. The organism was Gram-positive but
not acid fast.
It grew well on blood-agar and * nasgar,* as already stated.
When first isolated it grew scantily on agar, but subcultures were
not successful. It did not grow on glucose agar or maltose agar.
On potato it produced an abundant whiteish growth which developed
a brown or grey-brown efflorescence and stained the medium dark
brown. On blood serum it grew well, causing no liquefaction but
staining the medium dark brown. It did not grow on gelatin, nor
in ordinary broth and peptone water.
Its qualitative bio-chemical reactions were tested, and no change
was produced in any of the following:—Arabinose, Rhamnose
(iso-dulcite), Galactose, Glucose, Laevulose, Mannose, Lactose,
28s
Maltose, Saccharose, Raffinose, Amylum, Dextrin, Glycogen,
Inulin, Amygdalin, Helicin, Phlorrhizin, Salicin, Glycerol, Erythrol,
Adonitol, Dulcitol, Inosite, Mannitol, Sorbitol, and Litmus milk.
A guinea-pig inoculated intraperitoneally with an emulsion of a
culture appeared to be unaffected. This experiment, however, was
not made until seven months after the organism was isolated.
The organism showed the characters of a fungus of the Genus
Nocardia , but so far as we are able to ascertain, does not correspond
with any of the numerous species already described. Although it
was obtained in cultures made from blood aspirated from the heart,
it is not possible to say if the organism is pathogenic to man, but
the fact that its growth was almost restricted to potato and media
containing either blood serum or ascitic fluid is perhaps significant.
We propose for this fungus the name 'Nocardia cruoris.
SUPPLEMENTARY NOTE ON A CASE OF
BRONCHOMONILIASIS IN A NATIVE OF
THE GOLD COAST
A short time ago (1921) we described a fungus of the Genus
Monilia which we had isolated from a case of bronchomoniliasis at
Accra. The fungus belonged to the Tropicalis group of Castellani
and Chalmers, and closely resembled in its bio-chemical reactions
M . nivea , but we could not say at that time whether it was actually
the same- or distinct, because we were unable to test its action on
Raffinose.
We are now able to fill in this gap in our description, and to
state that neither acid nor gas is produced in Raffinose. The
fungus, at the time this reaction was tested, had been isolated and
maintained in cultures in the laboratory for about eight months, but
a re-examinalion of its other bio-chemical reactions showed that
they had not appreciably altered. The organism, therefore, cannot
be regarded as M . nivea , and as previously pointed out, it differs
from all the other members of the Tropicalis group; it must,
therefore, be regarded as a new species, for which we propose the
name Monilia accraensis.
286
In support of the view that the failure of the fungus to ferment
Raffinose was not due to the loss of this property after isolation
from the human host, it may be added that we have recently
obtained the same organism from the sputum of another native
patient at Accra, and that when tested immediately after isolation
it produced neither acid nor gas in this medium.
REFERENCE
Macfic, J. W. S., and Ingram, A. (1921). Bronchomomtiant Complicating Pulmonary
Tuberculosis in a native of the Gold Coast, West Africa. Annals of Trop. Med. &
Parasitol XV, pp. 53-58.
287
REPORT ON RAT-FLEA INVESTIGATION*
BY
R. NEWSTEAD, F.R.S.
AND
ALWEN M. EVANS, M.Sc.
(Received for publication 25 August , 1921)
INTRODUCTION
The investigation was carried out in conjunction with* Dr. E. W.
Hope, Medical Officer of Health for the City of Liverpool, and his
Assistant, Dr. W. Hanna, whose material assistance has made it
possible for us to carry it to a successful conclusion.
The object of the investigation was to determine the distribution
of the various species of fleas occurring on the rats in the Port and
City of Liverpool, with special reference to those species which are
responsible for the transmission of plague from rat to rat and from
rat to man. %
The period during which the investigation was carried out
extended from April 12th, 1920, to April 12th, 1921, rats being
examined daily five times each week, except during the month of
August and from December 22nd, 1920, to January 12th, 1921.
TECHNIQUE
As a preliminary to these investigations, field observations were
conducted as to the general methods adopted by the professional
rat catchers in the capture of the animals on board ship and in the
city.
It was then decided that the rats sent to the laboratory for
examination were to be taken alive from the traps, and placed singly
in strong calico bags. The bags were subsequently placed in a
lethal chamber, where they remained until the rats and the fleas
upon them were dead. This method ensured that all the fleas living
on each rat at the time of capture were secured. It was further
• Reprinted from The Annual Report of the Medical Officer of Health ( Liverpool) to the
Port Sanitary Authority , for the year 1920, by permission of Prof. £. W. Hope, M.D., with
additional charts, etc.
288
decided that, as far as possible, only rats caught singly in traps
were to be sent to the laboratory, because it was seen that, when a
number of rats are together in one trap, sweating is apt to occur,
with the result that the fleas leave the rats before the latter are
captured.
LABORATORY METHODS
The fleas were collected from each bag and rat, over a sheet of
white paper, and placed in a numbered watch-glass containing
glycerine. In this medium they were arranged in rows on a slide,
and identified by microscopical examination.
DEFINITION OF ZONES
• ZONE I.— Ships. The number of ships from which rats were
received was one hundred and twenty-five. The ports from which
the ships sailed are named in Table I ( vide infra).
ZONE II.— Docks, ETC. This Zone comprises that part of the
Portr of Liverpool lying west of a certain line, which defined as
accurately as possible the eastward boundary of the warehouse area
of the Port.
{a) Docks. This section of Zone II is limited to the wharves
themselves and the sheds actually situated on the Docks.
( b ) Warehouses. This section includes the rest of Zone II; the
majority of the buildings, from which rats were received, were
warehouses.
Zone III.— City. All of the City east of the line bounding
Zone II is included in this Zone.
THE SPECIES OF RATS
No attempt is made to separate the varieties of the species
Mus rattus and Mus norvegteus. Mus rattus and its varieties are
included under the heading * black rat/ and Mus norvegteus and
any varieties are referred to as * brown rats/
As will be seen from the following table, all the rats received from
ships were black. In Zone IlA the black rats outnumbered the brown
289
Table I.
Showing the number of rats received from ships sailing from the various ports.
Region
Port
No. of rats
Mediterranean .
Alexandria .
64
Constantinople ...
5
Costanza .
2
Genoa .
5
Salonika.
4
80
India, Ceylon, E. Indies, Burma
Bombay*.
5
Calcutta* .
18
Colombo* .
4
Rangoon* .
47
Java* .
1
75
East Africa... .
Beira* .
,
Mombasa*
3
Various Ports .
1
5
West Coast of Africa .
Various Ports .
59
West Coast of S. America .
Valparaiso .
5
Callao .
1 *3
Coronel.
. 18
Pisco* .
3
Talcahuano .
3
Various Ports .
33
75
Brazil .
Bahia*.
,
Pernambuco* .
18
Rio de Janeiro*.
13
Rio Grande .
5
37
Argentine.
Buenos Aires .
53
R. Plate.
*7
Rosario ...
*5
!
North America
Galveston
14
New Orleans .
1 4°
New York .
! 1
! 55
1
Within the Tropics.
290
rats by nearly five to one. In Zone IIb, the black rats were much
more numerous than the brown rats. In Zone III, however, the
position is more than reversed, the brown rats being nearly nine
times as numerous as the black rats.
Table II.
Showing the distribution of black and brown rats.
Zone
Black rats
Brown rats
Total
1 .
469
—
469
IIa .
24
5
2 9
IIb.
442
?74
716
Ill .
20
179
«99
Totals.
955
458
MI3
Table III.
Showing distribution of the various species of fleas.
Zone
Xenopsylla
cbeofis
CeratopbyUus
fasciatus
CeratopbyUus
hndiniensis
Leptopsylla
musculi
Ctenocepbalus
Car, is
Totals
I .
489
219
—
8
—
706
IIa.
10
75
—
9
—
94
IIb.
' 60*
1,5 IO
3
326
—
1,899
Ill.
3
320
12
10
1
346
Totals
562
:
2,124
\
*5
353
1
3.<>45
• This figure includes 56 specimens from No. 32 T-Street.
OBSERVATIONS ON THE VARIOUS SPECIES OF FLEAS FOUND
Xenopsylla cheopis , Rothsch.
The Indian Plague flea is ‘the common rat flea of the tropics*
(Rothschild, 1910). It is prevalent also in sub-tropical countries,
and ‘ is common during summer and autumn in some of the warmer
parts of the temperate zone, more especially in ports which have
maritime intercourse with the tropics* (Chick and Martin, 1911).
It has been demonstrated experimentally by Bacot (1914) that
temperatures below 40° F. are fatal to all but the imaginal stages
of this species. Its almost complete absence from more northern
latitudes is therefore attributed to the low winter temperatures which
prevail there. In England, X . cheopis has been recorded from a
brown rat at Plymouth (one specimen) (Rothschild, 1905); from a
plague-infected brown rat at Bristol (two specimens), 1916, taken in
a rag factory that was the seat of an outbreak of plague; and in
1911 from Guy’s Hospital, London, where the species had established
itself on a colony of brown rats. In the last case, it was found
that the infested rats were living in an artificially heated environ¬
ment, beneath the laboratory, and the presence of this, the first
flourishing colony of X. cheopis to be found in England, was
attributed to this fact.
Seasonal Variation (Chart II and Table VII). Zone I is the
only Zone from which a sufficient number of specimens of X. cheopis
have been found to allow of any deductions as to seasonal variation.
The curve, Chart II, has no very well marked characteristics; the
highest point reached, June, 1920, rises from low figures in both
May and July. The months September, October, November, show
a second but not so great elevation, falling to the lowest figure
in December.
In view of the prophetic statement put forward by Chick and
Martin (1911, p. 125), it is interesting to observe that we have found
a number of X. cheopis on the ship rats during every month in the
year, the lowest being 0 30 in December, 1921, and the highest
2*18 in June—not in September, as might have been expected.
Owing to the large numbers of Xenopsylla cheopis which occur
on ship rats, it would appear likely that this species might be present
on some of the rats found in the dock sheds. This was found to be
the case; out of the twenty-four rats received from Zone IlA, three
were found to carry X. cheopis. The data were as follows: —
Table IV.
No.
Date
Rat
Building
X. cheopis
Black
Brown
127
31.V.20
1
—
S. H-Dock
2
148
7.V1.20
1
—
S. C-Dock
7
192
23.vi.20
—
1
C-Dock
1
Total
2
1
10
292
In Zone IIb and Zone III, apart from the colony of X. cheopis
found in T- Street, which is dealt with in a separate section,
isolated specimens of this species were found in a few cases. The
data are as follows : —
Table V.
Zone IIb.
No.
,
Date
R/
IT
!
Building
X. cbeopis
Black
Brown
1019
21.1.21
—
1
14 C-Street
1
1127
10.it.21
—
1
29 C-Street
1
11 3 5
11 .ii.21
—
I
27 K-Street
1
1417
12.iv.21
—
I
M-Mills
1
Total
0
4
4
Zone III.
No.
Date
Rat
Building
X. cbeopis
Black
Brown
1068
3i.i.2i
—
1
5 B-Road
1
1401
8.iv.21
•
—
W-Village
:
2
Total
1
1
3
Ceratophyllus fasciatus , Bose.
This is the common rat flea of temperate countries. In the rural
districts of Suffolk and North Essex, Strickland and Merriman
(1913) found this species to comprise 60 per cent, of the flea
population of rats. Our figures show a much higher percentage of
C . fasciatus for the city of Liverpool:—78 per cent, in Zone IIb and
92 per cent, in Zone III.
Seasonal Variation (Chart III and Table VII). The curve of
frequency for Zone IIb reaches a markedly high level duving
the summer months, with elevations in May-June and again in
293
September. A distinct depression occurs in July. The temperature
curve (Chart I), on the other hand, is considerably higher in this
latter month than in May. In September it is lower than in July.
The frequency curve for Zone III is based on a much smaller number
of records than that for Zone IlB, but with one exception it possesses
the. same characteristics as the latter. The exception occurs in
January, when it rises to 4*11. This high figure is due to the
occurrence of two very heavily infested rats among a total of nine.
The average infestation of the other seven rats was only 07. It is
apparent that there are two periods of maximum prevalence of the
species, one in early summer, and a second in September. (The
latter may possibly begin during August.) These periods both
occur during the warmer half of the year, but the lack of detailed
correlation between the frequency and temperature curves makes it
clear that in Liverpool the prevailing atmospheric temperature has
not such a direct influence on the prevalence of the species, as
results obtained elsewhere in this country have suggested.
There is no correlation between the curves of average humidity
and frequency of C. fasciatus. It must be noted, However, that
the former is based on records taken in the open, and that the
atmospheric humidity probably differs considerably in the buildings,
sewers, etc., frequented by rats.
HOSTS. All the rats from Zone I were black, therefore no
comparison between the number of fleas found on the different species
of rats can be made for this Zone. In Zone II the infestation of
brown rats with C. fasciatus was rather heavier than that of the
black rats.
Leptofsylla musculi , Dug£s
This is a widely distributed species naturally parasitic on mice
(Mus muse ulus), but frequently found on rats where the latter inhabit
buildings, etc., frequented by mice. This species is a proved
potential carrier of plague bacillus, and ‘ a small percentage 9 will,
according to Bacot (1919), ‘bite man under certain conditions/
Strickland and Martin (1913) found it to comprise a very small
percentage of the flea fauna of rats in East Suffolk and North Essex,
only three specimens being taken in all. Bacot (1919), however,
considers that this species may be more prevalent in England than
294
this figure suggests. As the following figures show, this is the case
in the city of Liverpool: —
per cent. L. musculi .
Zone IlA . 97 per cent, of total.
Zone IIb ... ... ... 17*2 ,, ,,
Zone III . 2*8 ,, ,,
Seasonal Variation (Chart IV and Table VII). The curve
shows that the species is most prevalent in Zone II during the
months of June-September, after which the numbers fall away to
almost zero during December-April. This curve agrees largely with
that for C . fascia/us in Zone IIb, but the rise takes place a month
later, and the disappearance during the winter months is more
complete. As in the case of C . fasciatus, the curve rises to a high
level during September in spite of the drop in temperature during
this month.
HOSTS. The numbers of Z. musculi per rat were almost equal on
the two species, the averages were: —
Zone II.—Brown rat .,
Number of fleas per rat
o' 41
Black rat
• • »» » *
f t
C48
„ III.—Brown rat
M * 1
1 f
0‘05
Black rat
• * » » >»
> t
o - o5
Ceratophyllus londiniensis , Roths.
This species is said to be ‘ A rare Mediterranean species,
probably introduced by port rats, has occurred in the house mouse
(Mus musculus ), and possibly on the brown rat (Epimys norvegicus)
in London, Dover and Aberdeen.* (Rothschild, 1911.)
Its distribution in Liverpool is interesting; apart from two
isolated cases in Zone II, the records, four in all, came from a
certain area, where the species appears to have established itself.
Ctenocephalus cants , Curtis
The common dog flea. This species has frequently been recorded
from rats, sometimes in large numbers. It was therefore surprising
to find only one specimen, which occurred on a brown rat from
Zone III.
295
Ctenopthalmus agyrtes , Heller
Strickland and Merriman, and Nuttall and Strickland, working
in rural districts of East Anglia, found this species to comprise a
large proportion of the rat fleas in this part of England. Our
records, however, show a complete absence of this species.
Rothschild (1915) states that this species occurs on the ‘brown rat
. . . jiving in the fields, and on the bank vole . . . the common
shrew . . . and others.* It is not, therefore, surprising that it is
not found on rats in such an urban district as the City of Liverpool.
Further, ten other species of Pulicidae, which are recorded by the
above-named authors, from rats in East Anglia, and which are
naturally parasitic on birds and small wild mammals, are likewise
absent from our records.
PERMANENT BREEDING PLACES OF XBNOPSYLLA CHEOPIS
On the last day of January, 1921, a specimen of X. cheopis was
found on a brown rat from No. 32, T- Street. We requested
that more rats should be sent from this building, and altogether we
received during the first half of February thirteen brown rats from
which no fewer than fifty-six X. cheopis were taken. The records
were as follows: —
Table VI.
No.
Date
Brown
X . cheopis
C. fasciatus
1067
31.L21
1
1
—
1087
3.U.21
1
5
2
1094
4-ii.2i
1
5
7
io 95
4-ii 21
5
4
1096
4-ii.21
9
4
1097
4.U.21
1
7
2
1103
7-ii.ii
6
0
1104
7.ii.2i
1
2
8
1105
7-ii.z 1
1 1
6
4
1106
7-ii.21
1
1
0
1107
7-ii.21
3
4
1108
7.H.21
5
2
i *33
11.ii.21
i *
—
Total .
*3
56
37
Average X. cheopis per rat — 4*3.
296
There can be no doubt that here at T- Street is a second
‘ flourishing colony of X . cheopis in the British Islands.’ As in the
case of the first such colony to be discovered (Rothschild, 1911), its
existence is probably to be attributed to the fact that its hosts were
living in an artificially heated environment.
Beneath the middle of the roadway adjoining the premises
T- Street runs a steam culvert. The excavations, which it was
possible to make, revealed that the rat burrows lead in the direction
of the culvert. It was, therefore, thought probable, that the nests
Table VII.
Showing the average frequency during each month of the common species in the lones in
which they occur.
Date
Average
Mean
Tempera¬
ture
Average
Humidity
Zone I
Zone II ( b )
Zone III
Xe:iop-
sylla
cbeopis
Cerato-
pbyllus
fasciatus
Cerato-
pbyllus
fasciatus
Lcptop-
sylla
musculi
Ccrato-
pbyllus
fasciatus
1920—
April
46*3°
73
o *33
ns
12-66*
0*0
No rats
received
May
52*9°
74
0*59
0*90
5 ' 3 *
0*32
-
June
57 '*°
73
2* 18
0*40
3 -z8
1-84
3*27
July
57*2°
79
0-51
o'i6
2'70
0*89
1*07
August
5 *’ 4 °
80
fnvestigati
on suspend
ed.
September
55 ‘ 5 °
81
1*58
O’O
5*7
1-83
2*24
October ...
5 «‘i°
84
*75
0*22
1-82
046 j
1*09
November...
45-8°
83
i*i8
0*37
r6i
1
0*4* |
o-68
December
40-5°
86
0*30
0-09
I-2 9
0*06
0*58
1921—
January
46-0°
84
0-48
0*03
i-* 9
0*04
4** *•
February ...
4***°
84
°*93
075
**33
°‘°4
*•27
March
45 - 5 °
79
0-90
i
070
ro8
0*07
1*00
April
46 * 7 °
74
1*00
;
—
o*6i
0*09
0*40
# Based on a very small number of records.
297
of the rats were situated in the near neighbourhood of the steam
culvert, and as the temperature of the latter was found on 21.2.21
to be io 5°F., the area surrounding it for a considerable distance
must be maintained at a temperature higher than the normal.
Unfortunately it was impossible to trace the burrows far enough to
discern whether the nests were actually within the area influenced
by the steam culvert, and, therefore, the suggested explanation of
the presence of this colony of X. ckeopis can only be regarded as an
extremely probable one.
SUMMARY
1. Five species of fleas were found to occur on rats from
the ships, Port, and City of Liverpool. They were:— Xenopsylla
cheopis , Ceralopkyllus fascia/us, Leptopsylla ntusculi , Ceratophyllus
londiniensis , and Ctenocephalus cants .
2. Xenopsylla cheopis occurred freely on ship rats throughout
the whole period of the investigation. It was also found on three
rats from the dock sheds, and isolated specimens were found on four
rats from Zone IlB, and on two rats from Zone III. A permanent
breeding place of the species was discovered in certain premises in
Zone lib.
3. Ceratophyllus fasciatus was universally prevalent during the
whole course of the investigation. The number of fleas per rat was
greatest during the summer months, but the curve of frequency could
not be correlated in detail with that of the average temperature.
4. Leptopsylla musculi was most prevalent on rats from Zone II.
It occurred very rarely upon ship rats. Ceratophyllus londiniensis
was found rarely in Zones II and III, and of Ctenocephalus cants
one specimen was taken in Zone III.
298
Chabt I
60 - 0 ° F
60 0 ° F
40 0 ° F
April May Jane July Aug. Sept Oct. Not. Dec. Jen. Feb. Mar. April
Black line :—Average relative humidity. Dotted line :—Average temperature.
Chart II. Xenopiylla cheap is
Average number of fleas per rat in Zone I.
3 °°
April Mij Jane July Aug. Sept. Oct. Not. Deo. Jan. Feb. Mar. April
Average number of fleas per rat in Zone IIb.
REFERENCES
Bacot, A. W. (1919). Fleas and Rat? in relation to Plague. Journ. Rny. Sanit. Inst., Vol. XL,
No. 1, 1919, pp. 53-60.
-(1014). A study of the bionomics of the common rat fleas and other species associated
with human habitation, with special reference to the influence of temperature and
humidity at various periods of the life history of the insect. Journ. Hygiene , Plague
Supplt. Ill, Jan. 14, 1914.
Chick, H., and Martin, C. J. (1911). The fleas common on rats in different parts of the
world, and the readiness with which they bite man. Journ . Hygiene , Vol. XI, 1911,
pp. 122-136.
Martin, C. J., and Rowland (1914). Rat Plague in East Suffolk. Reports to Local Govern¬
ment Board on Public Health and medical subjects.
Nuttall, G. H., Strickland, C., and Meriuman, G. (1913). Observations on British Rat-
fleas. Parasitology , Vol. VI, No. 1, April 17, 1913, pp. 1-19.
Rothschud, N. C. (1905). Occurrence of Pule: e cbeopis Rothsch, at Plymouth. Ent. Mo.
Mag., Ser. 2, Vo!. XVI, p. 139.
- (1910)* A synopsis of the fleas found on Mus norvepicus dectmanus, Mus rattus
alexandrinus , and Mus musculus. Bull. Ent ., Re*. I, pp. 89-98.
- (1911)* A further note on Xenopsylla cbeopis, Rothsch. Ent. Mo. Mag., Ser. 2, Vol.
XXII, p.113.
- (1915). A synopsis of the British Siphonaptera. Ent. Mo. Mag., Ser. 3, Vol. I (
pp. 49-112, pis. VII, XIV.
- (1916). The occurrence of Xenopsylla cbeopis , Roths., in Bristol. Ent. Mo. Mag.,
Ser. 3, Vol. II, p. 279.
3oi
NATURAL ENEMIES OF
STEGOMYIA CALOPUS , Meigen
BY
C. J. YOUNG
From the Laboratory of the Liverpool School of Tropical
Medicine , Manaos
{Received for publication 25 ]uly t 1921)
Plates XIX and XX
During the examination of various collections of water in Mandos
for the larvae of Stegomyia calopus 9 their absence from many
apparently suitable breeding-places attracted attention. Larvae
were found capable of developing in water taken from some of these
places, except when certain insects were present. A collection of
various aquatic insects from ponds, igarapds, etc., was therefore
made and placed in jars of water together with the larvae of
S. calopus. Under these conditions the following were found to
destroy Stegomyia larvae with varying degrees of efficiency : larvae
of dragonflies, water bugs {Hemiptera) and their larvae, larvae of
water beetles, and two others not identified. Of these the larvae of
dragonflies and water bugs were found to be the most destructive to
the larvae of 5 . calopus in captivity.
The habits of 5 . calopus larvae, which constantly range about
in search of food, render them particularly liable to attack by
predaceous insects, as compared to certain Culex and Anopheles
larvae which may remain stationary at the surface for long periods.
The commonest mosquito larvae found in Mandos were those of
Stegomyia calopus and Culex fatigans \ these two species were
frequently discovered breeding together in wells, barrels and other
collections of water. The movements of both larvae and pupae of
C. fatigans differ from those of 5 . calopus in being much quicker,
thus rendering them less easily caught by predaceous insects. The
larvae of C. fatigans spend most of their time at the surface, but
302
descend occasionally. On the other hand, the larvae of 5 . calopus ,
although occasionally searching the surface film for food, while
taking in air, spend most of their time ranging over the bottom and
sides of their habitat and are thus particularly exposed to attack by
the dragonfly larva, which lies in wait at various depths below the
surface.
DRAGONFLY LARVAE
Dragonflies are present in Mandos throughout the year, there
being no cold season, and larvae were readily found at all times.
Although representatives of the AGRIONIDAE have been found in
Manaos, members of the AESCHNIDAE and LlBELLULlDAE have been
used exclusively in this work. Two species of each of these families
are shown (Plate XX, figs. 4 to 7). The larvae of the LlBELLULlDAE
(Plate XX, figs. 4 and 5) differ from those of the AeSCHNIDAE
(Plate XX, figs. 6 and 7) in having the abdomen shorter than the
hind legs. According to Miall (1912), the larvae feed on insects,
snails, tadpoles, and small fishes. In the laboratory they were
observed to attack other aquatic insects, including members of their
own species, water bugs and also tadpoles. I have never observed
them attempt to seize any object which was not moving. Larvae
have been observed resting on plants and on the sides and bottoms
of artificial ponds and fountains, igarapes and other natural
collections of water. They usually remain motionless and await the
approach of suitable prey, which they seize by suddenly shooting
out the labium. This is carried on a jointed arm which lies below
the head and thorax when not extended. The victim is then broken
up rapidly by the jaws and swallowed, only three or four seconds
being required for disposal of a Stegomyia larva. When the moving
object is out of range of the labium they sometimes spring towards
it by squirting water from the rectum. When disturbed they also
use this method of moving to a greater depth. Respiration is
carried out by drawing water into the rectum, but the more fully
developed larvae are said to be able to take in air through the
thoracic spiracles and, in the case of Aeschnid larvae, directly into
the rectum (Miall, 1912).
They are able to live in comparatively foul water. A dragonfly
303
deposited eggs in a barrel containing thick greenish water with
traces of oil on the surface and sides of the barrel and decomposing
vegetable matter at the bottom. Larvae of the type shown on
Plate XX, fig. 5, developed and lived for at least thirty-nine days,
by which time they had reached a size of 2 cms. in length. Observa¬
tions could not be continued. No dragonfly larvae were found in
water which did not receive rain.
One of the dragonfly larvae selected for experiment was identified
as Pantala fiavescens , F., by Dr. G. A. K. Marshall, who describes
it as a * migratory and almost cosmopolitan species.’ It was found
in several of the artificial ponds and fountains in public squares in
Manios (Plate XIX, figs. 1 and 2), in the igarap£s, and by Dr. R. M.
Gordon in a broken drainpipe under the pavement of one of
the public thoroughfares (Plate XX, fig. 3). In the pond shown
(Plate XIX, fig. 1) they were constantly present along with various
other species during the period of ten months that they were
observed. They destroyed S. calopus larvae and pupae of all sizes,
usually disposing of the larger ones first.
Six undetermined species of dragonfly larvae were also
experimented with, all of which proved destructive to the larvae of
S. calopus .
WATER BUGS, HBMIPTBRA
The distribution of the water bugs (Zaitha spp.) used in
the experiments is somewhat similar to that of the dragonfly larvae,
the former having been found occasionally in the pond shown in
Plate XIX, fig. 1, and in the igarap^s. They are not, however, so
plentiful, and difficulty was experienced in obtaining a supply at
times. They fed on aquatic and other insects, such as grasshoppers,
etc., when placed on the surface of the water, but only when living
and showing movement. They also attack their own species. They
suck the body juices of their victims, a specimen measuring 1*3 cms.
in length taking anything from five to fifteen minutes to dispose of
a fully grown 5 . calopus larva. The water bugs are thus consider¬
ably slower than the dragonfly larvae, but can continue feeding for
a longer period. They usually rest on plants or sides of ponds with
the posterior end of the body at the surface, the head downwards
3o 4
and the front pair of legs hanging free. When a suitable victim
approaches it springs on it by a quick movement of the hind legs
and seizes it with the fore legs which are adapted for clasping.
When the captured insect is large all the legs may be used for
gripping, and at least three mosquito larvae can be held at a time.
Both adult and larval water bugs fed freely on Culicine larvae.
FEEDING EXPERIMENTS
Experiments were carried out to test the relative efficiency of
dragonfly larvae and water bugs under artificial conditions.
Larvae of P. flavescens , measuring from 1*4 to 2*i cms., and
water bug larvae of ri to vg cms. in length, were used. Large
cylindrical glass jars containing two litres of tapwater were
employed, the depth of water in the jar being 10 cms. Twenty
fully grown mosquito larvae were used in each experiment and were
introduced before the insects to be tested, of which two were always
employed. The temperature of the water varied between 28*5° and
32 0 C. The jar was observed continuously for the first half hour,
and at three-quarter, one, one and a quarter, one and a half, two,
two and a half, three and five hours, and the number of larvae
destroyed during each period recorded. A control jar was used
containing twenty mosquito larvae similar to those used in the
experimental jar. Control larvae died in one experiment only, and
this is not included below. In the graphs shown in text-figs. 1
and 2 the ordinals represent larvae destroyed and the abscissae time
in hours. The curves in text-fig. 1 represent the averages of two
Fig. 1.
= a dragon fly larvae. -= a water bog larvae.-= 1 dragon fly + 1 water boglam.
S. = 20 larvae of S. calopus. C. *= 20 larvae of C. fatigatu.
3 °S
experiments in each case, but in fig. 2 each curve is constructed from
one experiment. It will be observed in text-fig. I that fastest time
was made by dragonfly larvae with S. calopus and second fastest
by a combination of dragonfly, water bug and Stegomyia larvae.
The other curves show only slight differences. In fig. 2 a mixture of
= 2 dragon fly larvae. -= 2 water bug larvae.-= 1 drag >n fly + 1 water bug larva.
io larvae of S. Cal opus.
io larvae of C. fatigans.
equal numbers of 5 . calopus and C. fatigans larvae were used, and
here a combination of dragonfly with water bug larvae was the most
effective. In the dotted curve the two dragonfly larvae destroyed all
the 5 . calopus in thtee minutes, but took five hours to the remaining
Culex larvae.
So far as rate of destruction was concerned, therefore, under
these artificial conditions, dragonfly larvae alone were the most
effective with S. calopus larvae, and a combination of water bug and
dragonfly larvae with those of C. fatigans.
As regards quantity, the largest number of fully grown 5 . calopus
larvae destroyed by a larva of P. flavescens in twenty-four hours
was one hundred and fifty-six. This larva during five days, being
fed on the first, third and fifth days only, consumed three hundred
and seventy-nine larvae. The largest number destroyed by a
water bug was one hundred and six in twenty-four hours, but only
half of these were killed for feeding purposes; a commoner figure is
about sixty. When about to moult these insects cease feeding, so
that the numbers consumed in twenty-four hours varied from o to
the maxima mentioned.
3°6
Some experiments with dragonfly larvae were carried out under
natural conditions.
An uncovered cement tank, measuring I metre square by 78 cms.
deep, in a backyard was filled with water to within 10 cms. of the
top. Between one thousand one hundred and one thousand two
hundred mosquito larvae in all stages of development were
introduced. The great majority of these were 5 . calopus , but a
number of C. fatigans were also included. Pupae and eggs were
also present. As a control, a glass jar was filled with water from
the tank and a few of both species of larvae added.
At zero (five hours later) both species of larvae could be detected
by a few moments inspection of the surface of the tank. Seven
larvae of P. ftavescens , varying in length from 23 to 1*3 cms., wer^
then introduced.
At sixteen hours both species of larvae could be detected.
At twenty-four hours no Stegomyia larvae could be found after
fifteen minutes search, but Culex larvae were still present.
At forty hours a few newly hatched Stegomyia and three Culex
larvae only were found. One dragonfly larva was found dead.
At forty-eight hours no larvae of any kind were found.
At fifty hours the tank was emptied by syphoning out the water
through several layers of gauze. No mosquito larvae were found.
In the control jar, which had been kept beside the tank
throughout the experiment, the larvae remained alive and active.
The temperature of the water in the tank varied between 29*5° and
33 5 ° C.
In the following experiment two barrels kept in an enclosure
behind the laboratory were used. Although frequently receiving
rain the water in these barrels was somewhat foul, particularly in
one which had recently contained oil. In size they measured
85 cms. by 56 cms. at the widest part and the depth of water in
each was 65 cms. Stegomyia calopus was breeding naturally in
both, large numbers of pupae and larvae in all stages of development
being present.
At zero (1 p.m.) five larve of P. flavescens were introduced into
one barrel.
At eighteen hours only small larvae were present.
At twenty-four hours no larvae were found.
J °7
During this period larvae and pupae continued to be present in
the control barrel to which the five dragonfly larvae were then
transferred.
At three hours pupae and larvae were still present.
At seventeen hours only small larvae were present.
At twenty-four hours a few small larvae were present.
At forty-two hours no larvae were found.
Thus two barrels were cleared of Stegomyia larvae in twenty-four
hours in one case, and in forty-two hours in the other, after the
introduction of five dragonfly larvae.
The tatter were allowed to remain in the last-mentioned barrel,
which contained the larvae of various other insects. During the
succeeding twenty-six days, five dragonflies hatched, no Stegomyia
larvae were observed, but on three occasions newly hatched Culex
larvae, which did not reach the pupal stage, were found. During
the last nineteen days of this period Stegomyia and Culex larvae
were present in the other barrel.
No Stegomyia larvae were found in Manaos in any water
containing dragonfly larvae.
A certain artificial well which contained dragonfly larvae and
was free from mosquito larvae, after being cleaned out was found to
be breeding Culex jatigans and Stegomyia calopus , but no dragonfly
larvae were discovered. Five weeks later Stegomyia larvae were
absent and dragonfly larvae were again found. A few Culex
jatigans larvae were also present.
THE DRAGONFLY AS AN ENEMY OF THE ADULT
MOSQUITO
It was thought probable that the adult dragonfly would prey
upon Stegomyia calopus if opportunity offered. In fine weather
dragonflies are to be seen in the open places and larger streets of
Manaos, and particularly around their breeding-places, as well as in
the forest. During dull weather they are not so active, resting most
of the time on the tops of shrubs and other objects, and rising only
in pursuit of insects appearing close at hand.
Both the dragonfly and Stegomyia calopus are, therefore, active
in Mandos in bright warm weather, but the mosquito is also active at
3°8
night. Except for a few attracted into Houses by lights, dragonflies
were not observed at night.
An attempt was made to see if they pursued S. calofus when
given the opportunity. It was found possible by moving slowly to
approach closely to the resting dragonfly without disturbing it, and
5 . calofus adults were then released from a glass tube at a distance
of about a metre from it. The dragonfly nearly always pursued and
caught the mosquito, usually returning to its perch consuming it.
It was not always possible, owing to the rapidity of its flight, to
observe whether the dragonfly actually caught the mosquito, but in
most cases it was seen to occur. Five S. calofus were released one
at a time near a resting dragonfly; four at least were caught,
and probably all five. The dragonfly was then caught, and
the contents of the alimentary canal teased up in saline and examined
microscopically. The mosquitoes were found to be broken up into
such small particles as to be unrecognisable, the only indication of
Stegomyia being the large number of flat scales present.
Five different species of dragonfly, varying in length from 3 cms.
to 6 cms., were tried against S. calopus , and all pursued and caught
the mosquito. The larger ones were also observed to prey upon
bluebottles and houseflies.
SUMMARY
Dragonflies and their larvae have been found to be destructive
to Stegomyia calofus and their larvae respectively. Several other
aquatic insects, including water bugs, have also been found to be
inimical to 5 . calofus larvae.
My thanks are due to the Director of the Laboratory,
Dr. Wolferstan Thomas, for assistance and advice, and to
Miss A. M. Evans and Dr. G. A. K. Marshall, Director of the
Imperial Bureau of Entomology, for identification of the insects
mentioned.
REFERENCE
Miall, L. C. (1912). Tbe Natural History of Aftsatic IntocUy Macmillan & Co.
3io
EXPLANATION OF PLATE XIX
Fig. i. Gardens in public square in Mangos. Habitat of larvae
of Pantala flavescens , F., and other dragonflies and
water bugs.
Fig. 2. Habitat of larvae of Pantala flavescens , F.
312
Fig- 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
EXPLANATION OF PLATE XX
Broken drain in street, Manaos. Habitat of larvae of
Pant ala flavescens, F.
Larva of Pantala flavescens. Actual length r8 cms.
Libellulid nymph. Actual length r8 cms.
Aeschnid larva. Actual length 2' I cms.
Aeschnid nymph. Actual length 47 cms.
.Innals Trop. Mtd. & ParasttolVol. XV
PLATE XX
C. Tinling < 5 ° Co. % LtdImp
3*3
WEST AFRICAN CERATOPOGONINAE
BY
A. INGRAM
AND
J. W. S. MACFIE
{Received for publication 24 August , 1921)
Plate XXI
This paper is a continuation of the series of papers on the
Ceratopogonine Midges of the Gold Coast already published in the
Annals of Tropical Medicine and Parasitology , and contains
descriptions of additional new species. .To some extent it is also
supplementary, since we are now able to fill in a number of gaps in
previous descriptions; for example, particulars are given of the early
stages and of the natural habitat of several of the species of
Culicoides and Dasyhelea , facts which are of interest, because they
point the way to means of control. It differs from the former
papers, however, in that a few species are described which were
collected, not in the Gold Coast, but in other parts of British
West Africa.
Many of the species belong to genera which were not considered
in the previous series; but the number of species referable to each
genus is small and insufficient to warrant the inclusion of detailed
generic descriptions. For the same reason, keys to the species found
in West Africa are not at present necessary. We have found it
difficult in some cases to appraise at their proper value the characters
employed by Kieffer in separating his smaller genera, and, where we
have found only slight divergencies to occur, we have been in doubt
whether it would be advisable, as would appear to be strictly
necessary, to erect new genera. We regret that in dealing with
these species we have not had the advantage of collaboration with
Mr. Carter, whose recent appointment as Malariologist in Ceylon
has unfortunately made it impossible for us to look to him for the
assistance he was ever so ready to give.
Some apology must be made for the figures which compare so
unfavourably with those in the previous papers. In the present
3i4
paper they are mere outlines traced with the aid of a camera lucida.
In figuring the hypopygium of the male we have not often
attempted to show it as a whole, but for the sake of clearness
and in consideration of our technical shortcomings, have drawn
the various organs more or less separated, showing them, unless
otherwise indicated, as they appear when seen in a ventral view.
Since our earlier notes on the bionomics of these insects were
published (1920) we have located another abundant source of
Ceratopogonine~midges, namely, the water lettuce Pist'ia stratiotes .
This weed, which is exceedingly abundant in rivers, swamps, pools
and lagoons in West Africa, and frequently covers large expanses
of water, has for many years been recognised as a troublesome
mosquito nursery and one very difficult to deal with, not only on
account of the association with it of Mansonioides africanus but also
because it provides protection between its leaves to the larvae of
several species of Anopheles ( e.gA. coslalis , A . mauritianus ,
A. nili ). The very small number of observations which we have
made hitherto prove beyond doubt that it is also a fruitful source of
midges, for we have reared from it Culicoides austeni , C. distincti -
pennis , Dasyhelea inconspicuosa , Prionognathus pseudomaculipennis ,
and the species described in the following pages under the names
Kempia ochrosoma 9 Eukraiohelea africana , E. versicolor , Probezzia
pistiae and P. stephensi. Large numbers of Kempia ochrosoma ,
Eukraiohelea africana and Probezzia pistiae were procured from this
source, so that it appears that these species, at any rate, are
peculiarly associated with the plant.
The types and co-types of the new species described have
been deposited in the museum of the Liverpool School of Tropical
Medicine.
Genus CULICOIDES, Latr.
Cultcoides austeni 9 Carter, Ingram and Macfie.
Five larvae of this species were found together with numerous
larvae of Ochlerotatus irritans in a sample of water from a crab-hole.
Three of the larvae were reared to the adult stage, the other two
were killed and preserved. The larval and pupal pelts of the
3*5
specimens reared through to the adult stage were recovered. The
following descriptions of the pupa and larva are based chi these
materials.
PUPA. Length about 27 mm. Respiratory trumpets short and
straight, tapering slightly towards the apex, raised on long stalks;
rather strongly chitinised, middle third covered with large squamose
spines; length of the trumpet about 015 mm., length of the stalk
about 0 05 mm. There are no knob-like processes, but the main
tracheal trunk gives off in its distal third a continuous row of about
nine short blunt processes. Cephalo-thorax : anterior marginal
tubercle double, highly chitinised, the inner portion large, conical,
and bearing a long strong bristle, the outer portion small, unarmed;
anterior dorsal very large, highly chitinised, irregularly conical,
bearing two stout bristles; dorso-lateral smaller, bearing a long and
a short hair; ventro-lateral an irregularly shaped tubercle bearing a
long and a short hair and an apparently unarmed socket; ventro-
median represented by a moderately long and a very minute hair.
External to the ventro-median tubercle and a little posterior to the
ventro-lateral is a small, unarmed, nipple-like tubercle which projects
prominently outwards. Dorsal tubercles small: anterior single,
bearing a minute blunt spine; posterior and lateral each bearing
a hair. Immediately in front of, and slightly external to, the
the anterior tubercle is a small tubercle which is apparently unarmed;
immediately behind, and slightly internal to the posterior, is a small
tubercle bearing a very minute spine. Postero-dorsal tubercle small,
bearing a hair, and behind it two unarmed sockets. Abdomen of
the usual form. Anal segment terminating in two sharply pointed,
divergent processes which are very highly chitinised, especially at
their tips. Tubercles on the abdominal segments small, strongly
chitinised; arrangement and armature as in C. accraensis.
LARVA. Length about 57 mm., greatest breadth about 0*3 mm.
Head : length about 0*2 mm., greatest breadth about 014 mm.
Eyes large, bilobed. Bristles small, apparently arranged as in
C. accraensis. Mental plate with a large, pointed central tooth.
Hypopharyngeal sclerite moderately chitinised, bearing on each side
usually eleven pointed, finger-like processes which are nearly equal
in size, excepting the fifth from the inner margin, which is slightly
larger than the others. Mandibles well chitinised, pointed, with a
well-developed central tooth. . Body : hairs minute,-terminal hairs
on the anal segment small; anal gills of the usual form.
Gold COAST: Accra, April, 1921, reared from mud from pools
and puddles near the station for the Weshiang Line (PI. XXI,
fig. 2 )> June, 1921, larvae found in water from a crab-hole.
Christiansborg, July, 1921, reared from plants of the water weed
Pistia stratiotes. Oblogo, June, 1921, reared from banana fibre.
Culicoides distinctipennis, Aust.
Although numerous specimens of this species have been collected
at Accra, it is only recently that we have reared it from the early
stages. In the samples from which it was reared were also the
early stages of several other species of Culicoides , and we were
unable to identify with certainty the larvae. The pupa was, however,
obtained, and is briefly described here.
PUPA. Description based on a single pelt from which a male
had emerged. Length 17 mm. Respiratory trumpets , raised on
moderately long stalks; length of the trumpet about 0 2 mm., length
of the stalk about 0 04 mm. Distal extremity somewhat darkened,
middle portion covered with squamose spines, proximal two-thirds
bearing three small knob-like processes. Main tracheal trunk
terminating distally in a fan-like arrangement of seven short, blunt
processes. Cephalo-thorax dark, operculum rather sparsely clothed
with large, dark, squamose spines. Anterior marginal tubercle
small, bearing a rather short, stout spine; anterior dorsal bearing a
short, stout spine and a minute spine; dorso-lateral, small, bearing
a hair and a short spine; ventro-lateral, small, bearing two hairs,
one of which is quite short and spine-like; ventro-median represented
by a moderately long and a very small hair. Dorsal tubercles:
anterior double, the two halves separated, the one situated anterior
and slightly external to the other, each bearing a short, stout, dark-
coloured spine; posterior, small and flat, bearing a minute spine;
lateral, small, bearing a hair. Posterior to the lateral of the dorsal
tubercles is an unarmed socket, and on the dorsum are several
ill-defined darkened patches resembling flat, unarmed tubercles.
Postero-dorsal tubercle small, bearing a hair and an unarmed
socket. Abdomen of the usual form. Anal segment terminating
in two sharply pointed, somewhat divergent, dark-tipped processes
which are not closely covered by squamose spines* Dorsal, ventro¬
lateral, and ventral tubercles as in C. accraensis .
Gold COAST: Accra, February to April, 1921; numerous
specimens reared from moist soil and mud taken from the margins
of pools and puddles near to the railway station on the Weshiang
Line (Plate XXI, fig. 2). Oblogo, March, 1921; reared from plants
of Pistia stratiotes taken from the river Densu (Plate XXI, fig. 1).
Culicoides eriodendroni , C., I. and M.
In a previous paper a description was given of the female of this
species. We are now in a position to add certain points in regard
to the male.
Measurements (average of two).
Length of body* .‘ . 1*3 mm.
Length of wing .. . . romm.
Greatest breadth of wing.0*3 mm.
In general, the description of the female applies also to the
male, but the following points may be especially mentioned. Eyes
separated. Antenna: last three flagellum segments sub-equal, the
fifteenth being, however, slightly the longest and the fourteenth
slightly the shortest. Scutellum bearing two central and two lateral
bristles, and a few (six) short hairs. Wing: pale spot covering the
anterior cross-vein rather diffuse and spreading almost to the costa,
pale spots at the apex of the wing and along its posterior border
almost or entirely absent; decumbent hairs scanty, but more than
a single row in the basal portion of the wing between the fourth and
fifth veins.
Hypopycium (fig. 1). Ninth segment : tergite, posterior margin
slightly notched in the middle and ending on each side in a well-
developed conical process; stemite, deeply excavated. Forceps :
side-pieces rather long and narrow, covered with relatively short
hairs; claspers with a broad, hairy, basal portion, constricting
abruptly to meet the terminal portion, which is of the usual form.
Harfes (fig. 1 d ): moderately chitinised, somewhat strap-like,
# In all case* taken from the anterior margin of the thorax to the tip of the abdomen of
specimens mounted in carbolic.
318
tapering only slightly, with a short piece at the distal end bent
ventrally or ventro-laterally like the end of a cleek. Aedoeagus
(fig. i b ): form rather unusual; stem very short, highly chitinised,
in ventral view shaped like the letter T; limbs long, moderately
highly chitinised; ventral wall chitinised for only a short distance
from the apex of the arch, membranous portion not spiculated.
Fig. i. CulicouUs rriodtndroni , C. I. & M., outlines of male hypopygium, ventral
view, a —forceps and harpes; k —aedoeagus.
Gold Coast: Nsawam, August and October, 1920; reared from
larvae obtained from rot-holes in the stuiqp of a silk-cotton tree and
of another tree.
Culicoides grahanti, Aust.
Several specimens of this species were reared from material taken
from the base of a banana stump. The pupal pelt of one female
was recovered, and as it was in some respects peculiar and showed
several characteristic features, a brief description of it is given.
PUPA. Length r6 mm., moderately well chitinised. Respira¬
tory trumpets long and curved; length about 0'i7mm. Stalk or
pedicle moderately long. The trumpets are of almost uniform
width throughout their length, and are irregularly ringed. The
main tracheal trunk gives off during its course through the trumpet
a few short lateral branches which terminate in small tubercles
scarcely projecting above the general surface level, and ends distally
in a fan-like arrangement of the usual form. Cephalo-thorax:
3 l 9
anterior marginal tuberde large, covered with squamfose spines,
bilobed, and formed of an inner rounded portion and an outer
conical portion, the latter bearing at its apex a long, strong bristle;
anterior dorsal, double, each part bearing a long, stout bristle;
anterior dorso-lateral, an irregular tubercle bearing two hairs;
ventro-lateral, an irregular tubercle bearing two delicate hairs,
ventro-median, apparently absent. The operculum is sparsely
covered with squamose spines, most strongly developed along its
periphery; near its posterior margin, in the middle line, is a small
elevation or tubercle covered with rather coarse squamose spines, but
otherwise unarmed. Dorsal tubercles small: anterior, double,
bearing a delicate hair internally and a small spine externally';
posterior, almost obsolete, bearing a small spine; lateral, bearing a
hair. Postero-dorsal tubercle almost obsolete, bearing a delicate
hair. Abdomen : anal segment bearing two transverse rows of
relatively large spicules, the one near the anterior margin and the
other about the middle of the segment; these spicules are developed
most highly on the dorsal and lateral aspects. A group of similar
spicules is present dorsally at the roots of the terminal processes.
The terminal processes are short and pointed at their tips, they
diverge almost at right angles, and their ends are turned dorsally.
Tubercles on the abdominal segments poorly developed, and shaped
like large spines. Dorsal tubercles: antero-submarginal, the inner
bearing a short spine and the outer a longer hair; postero-marginal,
only a single tubercle present, situated posterior to the outer
antero-submarginal tubercle, bearing a short spine. Ventro-lateral
tubercles small, but little larger than the dorsal and ventral: antero-
submarginal bearing a short spine; postero-marginal, the middle
one bearing a hair, the other two short spines. Ventral tubercles:
the middle one bearing a hair, the outer and inner each a short
spine. Each abdominal segment (excluding the last, which has
been referred to already) bears a transverse row of relatively well-
developed spicules near its anterior margin; these spicules are
most highly developed and most numerous on the more posterior
segments.
GOLD COAST: Nsawam, 24th July, 1920; reared from material
taken from the base of a banana stump.
320
Culicotdes neavei , Aust.
Several specimens of this species were reared from soft mud
taken from the edges of pools at Accra. The larvae, together with
those of several other species of Culicotdes , frequented the mud
which was very soft and almost semi-fluid. They were of the usual
form, but we are unable to give details of their structure because,
although we reared one adult from a larva isolated in a small tube,
we did not succeed in recovering the larval pelt, and were, therefore,
unable to distinguish amongst the numerous larvae in the
sample those belonging to this particular species. The pupae also
frequented the mud : the description which follows is based on an
examination of the pelts of two pupae which had been isolated
singly, and from which adult insects were procured.
PUPA. Length about i g mm. Operculum densely covered with
dark brown squamose spines. Respiratory trumpets similar to those
of Culicoides inornatipennis , short and raised on relatively long
stalks; length of the trumpet about 019 mm., length of the stalk
about 0 05 mm. The trumpet is infuscated at its distal end, and
also, slightly, at its base; it bears three or four quite small knob-like
processes, the most distal of which is situated rather far anteriorly.
The main tracheal trunk terminates distally in a hand-like group of
about six short blunt processes. Cephalo-thorax : anterior marginal
tubercle small, dark coloured, conical, bearing a relatively long
stout spine which is directed ventrally, anterior dorsal well
developed, conical, bearing a short stout spine and a minute spine;
dorso-lateral prominent, bearing a hair and a short spine; ventro¬
lateral a rounded hump, bearing a short and a moderately long
htfir; ventro-median small, bearing a moderately long and ^ short
hair. External to the ventro-median tubercle, and a little posterior
to the ventro-lateral, is a small nipple-like tubercle, apparently
unarmed. Just in front of and internal to the base of the stalk of
the trumpet is a small hair. Dorsal tubercles: anterior double, the
two parts being separate and well developed but not large knobs,
the one situated anterior to the other, each bearing a short, stout,
dark-coloured spine; posterior, poorly developed, bearing a minute
spine; lateral, feebly developed, bearing a hair. In front of the
anterior tubercle, and a little external to it, is an inconspicuous
unarmed tubercle; posterior to the lateral tubercle is a socket-like
321
mark, apparently unarmed, and there is another similar mark
situated more posteriorly. Postero-dorsal tubercle small, bearing a
hair and two apparently unarmed, socket-like marks. Abdomen :
anal segment with sharply-pointed terminal processes infuscated at
their tips. Dorsal tubercles of the normal form : antero-submarginal,
the inner bearing a short spine, and the outer a hair; postero-
marginal, five in number, the outermost bearing a hair, the next a
short spine, the innermost a minute spine, and the other two
apparently unarmed. Lateral tubercles larger than the others and
each with two sharp points, between which are the hairs or spines:
antero-submarginal, bearing a spine; postero-marginal, the npddle
one bearing a hair, the other two, spines. Traces are visible of a
second, rudimentary, antero-submarginal tubercle in a more dorso¬
lateral position. Ventral tubercles of normal form : the middle one
bearing a hair, the other two, spines.
GOLD Coast: Accra, February, 1921; reared from soft mud
taken from the edges of pools and puddles near the station for
the Weshiang railway (PI. XXI, fig. 2). From the same material
were reared also Culicoides austeni , C. distinctipennis , C. similis ,
C. schultzei and Stilobezzia spirogyrae.
Culicoides similis , C., I. and M.
At the time wfien this species was described (1920), the early
stages were not known. They have since been collected, and are
here briefly described.
Pupa, Length about rg mm. Operculum densely covered with
dark brown squamose spines. Respiratory trumpets short and
raised on rather long stalks; length of the trumpet about 019 mm.,
length of the stalk about 0 03 mm. The trumpet bears on 'its
proximal half three or four small knob-like processes which are
infuscated. The distal end of the trumpet is dark brown : in it the
main tracheal trunk terminates in a fan-like group of short, blunt
processes. Cephalo-thorax : anterior marginal tubercle dark brown,
rather small, conical, bearing a relatively long, stout, dark-coloured
spine; anterior dorsal prominent, irregularly conical, bearing a stout
spine and a minute spine; dorso-lateral small, bearing a hair
and a minute spine; ventro-lateral a rounded hump, bearing a
small and a minute hair; ventro-median represented by two hairs,
322
one minute. Dorsal tubercles: anterior usually double (in one
specimen single on one side), the two parts contiguous or separated
but almost side by side, well developed but not large knobs, each
bearing a short, stout, dark-coloured spine; posterior, poorly
developed, bearing a minute spine; lateral, poorly developed,
bearing a hair. In front of the anterior tubercle, and a little external
to it, is an unarmed tubercle; posterior to the lateral tubercle is a
socket-like mark, apparently unarmed, and there are usually two
similar marks situated more posteriorly. Postero-dorsal tubercle
small, bearing a small hair and one or two apparently unarmed,
socket-like marks. Abdomen : anal segment with sharply-pointed
terminal processes infuscated at their tips; in the middle line,
dorsally and posteriorly, is a small elevation covered with dark
squamose spines. Dorsal tubercles of the normal form usually, but
on some segments the outer two postero-marginal tubercles tend
towards the form of the lateral tubercles: antero-submarginal, the
inner bearing a short spine, and the outer a hair; postero-marginal,
five in number, the outermost bearing a hair, the next a spine, the
innermost a, minute spine, and the other two apparently unarmed.
Lateral tubercles larger than the others, and each with two sharp
points between which arises the hair or spine: antero-submarginal,
bearing a spine; postero-marginal, the middle one bearing a
hair, the other two, spines. Traces are visible of a second,
rudimentary, antero-submarginal tubercle in a more dorso-lateral
position. Ventral tubercles of normal form : the middle one bearing
a hair, the other two, spines.
LARVA. Although we did not succeed in recovering the larval
pelt of any individual specimen isolated and reared through from
the larval stage, we secured from the materials collected at Accra a
number of larvae which we believe to be those of Culicoides similis.
The larvae were found, together with pupae of C. similis , in the soft,
semi-fluid mud in the specimen jar at a time when this species was
the only one emerging from the sample. Moreover, in three of the
larvae, apparently almost ready to pupate, the pupal structures,
including the respiratory trumpets and many of the cephalo-thoracic
and abdominal tubercles, were clearly visible through the cuticle,
and appeared to be identical with those of the pupa of C. similis .
3 2 3
It is of interest to note the situations in which certain of the
pupal structures were seen in the larvae. The trumpets lay in the
first body segment, their proximal ends situated dorsally and
laterally at the posterior end of the segment, and their free ends
situated ventrally on each side of the middle line a little posterior
to the head. The dorsal tubercles of the cephalo-thorax were
situated dorsally at the anterior end of the second body segment.
The terminal processes of the anal segment were turned so that they
projected anteriorly, one on each side, with their tips directed
dorsally and situated at the anterior margin of the segment.
The following description of the larva is based on the examina¬
tion of the three larvae alluded to as being apparently ready
to pupate.
Length about 3*5 mm. when fully grown, greatest breadth about
002 mm. Head , length about 01 3 mm., greatest breadth about
0 08 mm. Eyes small. Bristles mostly small; on the ventral surface
one pair, admedian, a little posterior to the hypopharyngeal
sclerite, two pairs, admedian, almost contiguous, anterior to the
hypopharyngeal sclerite, and one pair ventro-lateral; on the lateral
surface two pairs, the one anterior and the other central; on
the dorsal surface one pair admedian, anterior, two pairs sub-
central, slightly separated, dorso-sublateral, two pairs, almost
contiguous, posterior, dorso-lateral. Palpi and antennae well
developed. Labium broad, blunt, dark coloured, apparently without
teeth. Posterior margin of hypopharyngeal sclerite bearing on each
side seven or eight sharply pointed, graded, teeth, the middle
ones being the longest; these teeth are not very highly chitinised.
Mandibles simple, pointed, without teeth. Body : appearing almost
hairless, but actually bearing a few very small hairs; terminal hairs
on the anal segment very small; anal gills of the usual form, deeply
cleft distally into two pointed processes.
GOLD COAST: Accra, February to April, 1921; numerous
specimens of both sexes reared from soft mud taken from the edges
of pools and puddles near the station for the Weshiang railway
(PI. XXI, fig. 2). Oblogo, February, 1921 ; a few specimens reared
from sandy mud taken from the washing-place in the river Densu.
3 H
Culicoides corsoni , sp.n.
This insect, of which we possess at present only a single male,
resembles in wing markings Culicoides similis and C. citroneus.
From the former it differs in having no pale area covering the
middle of the lower ramus of the fourth vein and in the whole of the
cross-vein being enveloped in a.pale area; from the latter in having
only a single pale area between the branches of the fifth vein, the
small pale spot in the angle being absent.
Measurements.
Length of body (one male) .ro mm.
Length of wing . .o*8 mm.
Greatest breadth of wing. ... .0*3 mm.
Head : occiput dark brown. Eyes narrowly separated. Proboscis
and palpi dark brown; the latter with the third segment somewhat
inflated, especially in its basal two-thirds. Antennae : infuscated;
the thirteenth segment slightly longer than the terminal segment;
the plumes only moderately developed. Thorax dark brown, with
paler brown markings. Owing to the fact that the species was not
recognised until the hypopygium had been examined in carbolic,
exact details of the thoracic adornment cannot be given, but there
appeared to be three large paler areas on each side of the anterior
two-thirds of the dorsum, one in front and two behind. Scutellum
yellowish-brown, somewhat darker at the sides; bearing one central
and two lateral bristles, and a few (three) short hairs. Post-
scutellum dark brown. Pleurae dark brown. Wings with pale
markings resembling those of Culicoides citroneus . The pale spot
in the middle of the anterior border covers the tip of the distal cell;
there is no pale spot covering the middle of the lower ramus of the
fourth vein, and the pale spots in the neighbourhood of the
bifurcation of the fifth vein are arranged as in C . similis.
Decumbent hairs scanty, not extending beyond the middle of the
wing, and none being present in the anal angle, between the rami
of the fifth vein, or at the base between the fourth and fifth veins.
Halteres with creamy-yellow knobs. Legs infuscated, knees scarcely
darker; tibiae with narrow pale bands basally and less distinct pale
bands on the femora apically. Abdomen dark brown.
Hypopygium. Ninth segment: tergite broad, bearing only a few
stout hairs, the most notable of which are four arranged in a
3 2 5
transverse row near the posterior margin; posterior margin notched
in the middle, and bearing at each lateral angle a long, slender,
finger-like process about five times as long as it is broad at its
base; apical lobe-like processes well developed. Sternite deeply
excavated centrally. Forceps of the normal form; claspers highly
chitinised, with rather blunt ends. Harpes resembling those of
C. grahami , but with shorter distal portions. Aedoeagtis Y-shaped,
the limbs and the proximal end of the stem highly chitinised, the
distal end of the stem more delicate and ending in a broad lip.
The ventral wall of the aedoeagus is highly chitinised from the apex
of the arch to about the middle of the limbs, but is not extended
anteriorly and centrally as a spine; the membrane joining the
aedoeagus to the ninth sternite is without spicules. The hypopygium
resembles that of C. c it rone us, but the following differences, amongst
others, may be noted. The long hairs at the posterior end of the
ninth tetgite are less numerous than in C. citroneus , the lateral
finger-like processes more slender, and more pointed; the claspers
are* more highly chitinised; the harpes resemble more closely those
of C . grahami than those of C. citroneus ; and the proximal ends of
the limbs of the aedoeagus are inverted.
Gold Coast: Koforidua, April, 1921 (Dr. J. F. Corson); one
male taken in a bungalow, on the wall near a lamp. We have
pleasure in dedicating this species to the collector, Dr. J. F. Corson.
Culicoides nigeriae , sp. n.
Measurements.
Length of body (three females) . ... V2 mm.
Length of wing .0*9 mm.
Greatest breadth of wing.0*4 mm.
Head dark brown. Eyes separated, internal chitinous thickening
well developed; the -eyes are more widely separated than in
C. inornatipennis , the length of the internal chitinous thickening
being about 20 p instead of about 13 fi. Proboscis dark brown.
Palpi brown, third segment moderately inflated. Antenna dark
brown; segments four to ten from once to once and a half as long as
broad. Thorax uniformly dark brown, pollinose, sparsely clothed
with dark brown hairs. Scutellum uniformly dark brown; bearing
two admedian and two lateral bristles, and a few short hairs. Post-
326
scutellum dark brown. Pleurae dark brown. Wings unspotted;
distribution of decumbent hairs similar to that in C. inornatipennis ,
but wings rather less hairy. Halteres darkish brown. Legs
rather dark brown, almost unicolourous. Abdomen dark brown.
Spermathecae two, dark brown and very highly chitinised, sub-
spherical to oval, measuring about 35/“ by 30 p on an average; only
the very commencement of the duct is chitinised.
NIGERIA (Northern provinces): Gimi, Zaria Province, 27th
October, 1920 (Dr. W. B. Johnson); ‘collected whilst biting.’
Duchi-n-wai, about forty miles from Zaria, at an elevation of
2,000 ft. approx., 25th November, 1920 (Mr. L. E. B. Pearse), taken
in the act of biting. Ilorin Province, Kaduna river, 21st December,
1920 and 23rd December, 1920; Pategi, 24th December, 1920
(Dr. J. R. C. Stephens); some of the specimens taken on the arm,
and undoubtedly biting. Several specimens obtained from each
locality, all females.
This species, besides being a much darker brown insect, may be
distinguished from C. inornatipennis by the following points amongst
others: the absence of the characteristic thoracic; adornment, the
colour of the scutellum and the colour of the halteres.
Culicoides inornatipennis , C., I. and M., var. rutilus , var. nov.
A small rufous variety of C . inornatipennis , C., I. and M.
Length less than 1 mm., usually about o*8 mm.; length of wing
about 07 mm., and greatest breadth about 0 3 mm. Hypopygium
of the male and spermathecae of the female as in C. inornatipennis ;
other morphological characters also similar. Fourth segment of the
palp very small, about half the length of the fifth; scutellum bearing
three or four stout bristles, one or two central or admedian and two
lateral, and a variable number (about half a dozen usually) of smaller
hairs.
Colouration notably different from that of C. inornatipennis.
Head dark brown. Proboscis and palpi pale brown. Antenna
with a dark brown torus and lighter brown flagellum segments,
bearing pale brown hairs. Thorax unicolorous, light brown or
almost golden-brown. Scutellum light brown ; post-scutellum darker
but not very dark brown. Pleurae brownish-yellow. Halteres pale
3 2 7
yellowish-brown. Legs pale brown, almost unicolorous, but with
knee spots a* slightly deeper yellow-brown. Abdomen dark brown.
PUPA. Two pupal pelts were examined, and were found to be
indistinguishable from those of C. inornatipennis. They were very
small, length about 1*3 mm.
GOLD COAST: Nsawam, May to August, 1920; numerous
specimens reared from rotting fibrous material taken from the bases
of banana stumps. October, 1920; one specimen reared from
material taken from a rot-hole in a silk-cotton tree.
f Genus DASYHELEA , Kieff.
Dasyhelea fuscipleuris , C., I and M.
In a previous paper, Part IV of this study (1921), this species was
described. At that time we possessed only two females, taken in
buildings, and had identified neither the early stages nor the
habitat. Recently we have reared a large number of specimens, and
are able to supplement our previous description by giving an account
of the pupa. It is interesting to note that all the specimens reared
were females and to compare this observation with those previously
made on C. clarkei and C . eriodendroni (1920).
PUPA. Length about 2*1 mm., delicately chitinised and rather
slender. Respiratory trumpets very long and slender, length about
o*6 mm., breadth about 19^, raised on small tubercles; the main
tracheal trunk is narrow, gives off throughout its length, beginning
at its very base, numerous (about fifteen) quite short lateral branches,
and ends distally in a cluster of about six short processes. Cephalo -
thorax not very strongly chitinised, operculum coarsely shagreened.
Anterior marginal tubercle large, conical, bearing a small spine-like
hair; dorso-lateral irregularly shaped, bearing two delicate hairs;
anterior dorso-median small, bearing two very short, straight hairs;
ventro-lateral almost obsolete, bearing two minute hairs, ventro-
median represented by a small hair. Dorsum of the thoracic region
not infuscated, without tubercles, but with several brownish macules.
Abdomen feebly chitinised, of the usual form; tubercles small and
poorly chitinised, terminal processes widely divergent.
Larva. The larva is of the usual form. Only a single pelt
actually correlated with an adult was obtained, and in it no
328
characteristic points could be made out. Unfortunately, it was a
practical impossiblity to isolate the larvae of this insect from the
materials in which they were living, because the sample contained
also numerous other species ( Culicoides schultzei , C. similis ;
C. ausleni, C. dislinctipennis , Stilobezzia spirogyrae and Dasyhelea
inconspicuosa).
The single larva reared through to the adult stage was isolated
in a small tube containing nothing but a little damp filter paper.
The larva buried itself immediately but a day or two later the pupa
was observed worming its way to the surface through the mass of
filter-paper. The trumpets could not be distinguished with the aid
of a hand lens. Two days later a female C. fuscipleuris emerged.
The pupal pelt was found on the side of the tube half an inch above
the filter-paper, a situation which the pupa must have reached by its
own unaided efforts. The larval pelt was found embedded in the
filter paper near the spot where the pupa was first observed.
GOLD Coast: Accra, Apfil, 1921; numerous specimens (all
females) reared from mud taken from the edges of pools and puddles
near the station for the Weshiang railway line (PI. XXI, fig. 2).
Dasyhelea nigricans , C., I. and M.
This species was originally described from two males taken in
the laboratory at Accra. The halteres of these specimens had
yellowish-brown knobs and dark brown stems. More recently we
have received from Dr. W. B. Johnson a number of specimens of a
species of Dasyhelea taken at Kaduna, Nigeria, in August, 1920,
which resembles in every respect Dasyhelea nigricans , excepting
that in some the halteres are white, and in others yellow. We
conclude, therefore, that in this species the colour of the halteres
must be variable.
A few of the specimens collected by Dr. Johnson at Kaduna were
females, and we are, therefore, able to supplement our previous
account by describing the characters of this sex.
Measurements. Female.
Length of body .. .. 1*3 mxn *
Length of wing ... -. ro mm.
Greatest breadth of wing.0*35 mm.
3 2 9
The female resembles the male in most respects, but the following
points, including the more important differences, may be mentioned.
Head : eyes separated. Antennae : hairs dark brown, short and
scanty, segments of the flagellum gradually elongating from base to
apex in a continuous series, that is without an abrupt change
of form between the tenth and eleventh segments; segments four to
ten from once- to nearly twice as long as broad, segments eleven to
fifteen from twice to a little over three times as long as broad, the
last segment ending in a stylet; long spines present on all the
segments excepting the last five. Thorax : scutellum almost entirely
yellowish-brown; armature of bristles and hairs as in the male.
Wings rather densely .hairy, the hairs extending basally beyond the
cross-vein; bifurcation of the fifth vein at about the same level as
the termination of the costa; cell formed by the first and third veins
at their junction with the costa larger than in the male. Halteres
yellow. Legs : claws small, simple, equal. Abdomen clothed with
yellowish-brown hairs; spermatheca single, highly chitinised, sub-
spherical (diameter about 42/1), the commencement of the duct
chitinised for a considerable distance, about 15/n, most strongly at
its end of origin from the spermatheca.
NIGERIA (Northern provinces) : Kaduna, August, 1920 (Dr. W. B.
Johnson).
Dasyhelea nigeriae , sp. n.
Measurements. Male. Female.
Length of body . i*i mm. ri mm.
Length of wing .o # 8 mm. o # 8 mm.
Greatest breadth of wing.0*3 mm. 0*3 mm.
Head dark brown. Eyes separated in both sexes. Clypeus and
proboscis dark brown. Palpi dark brown; third segment inflated,
in the male especially basally, about the same length as the second;
fourth segment short, about half the length of the third. Antennae
unusually short, dark brown, with dark brown hairs; in the female
segments four to fourteen gradually lengthening towards the apex,
the length varying from two-thirds to once the breadth, the last
segment rather larger and longer and ending in a conical tip, but
without a stylet, in the male segments four to eleven short, those at
the base broader than long and those at the apex sub-spherical,
330
length varying from about two-thirds to once the breadth, last four
segments sub-equal, about four times as long as broad, not binodose
but exhibiting the usual sculpturing (compare D. flava ), the last
segment longer and stouter but without a stylet. Thorax uniformly
dark brown, with paler, somewhat yellowish, humeral patches.
Scutellum dark brown, bearing in both sexes one central, two
admedian, and two lateral bristles and no short hairs. Post-
scutellum dark brown. Pleurae dark brown. Wings hyaline,
without spots; clothed with decumbent hairs which do not extend
basally beyond the cross-vein. Venation as in D. flava ; interspace
very small in the male, more well developed in the female.
Halteres with dusky orange-coloured knobs and infuscated stems.
Legs brown, femora and tibiae dark brown, but lighter coloured in
the male than in the female. Claws short, equal, simple; in the male
bifid at the tips. Empodium rudimentary, but in comparison with
the size of the claws appearing to be larger than usual. Abdomen
dark brown, venter slightly paler than the dorsum. Spermatheca
single, highly chitinised, pyriform (about 42/* by 38/1); chitinised
part at the commencement of the duct conical, about 11 fi long.
Hypopygium (fig. 2). Ninth segment : tergite tapering slightly
posteriorly, sparsely clothed with strong hairs, posterior margin not
• 'W*
Fig. 2. Dasybelea nigeriae , sp.n., outlines of male hypopygium, ventral view.
a —ninth segment and forceps ; £--harpes; c —aedoeagus. ( x 375 circa.)
33 1
notched, and without finger-like lateral processes; stemite prolonged
posteriorly on each side of the middle line into a highly chitinised,
pointed, finger-like process. Forceps: side-pieces short and
stout, broader apically than basally, rather scantily clothed with
modefately long hairs } and bearing on the inner aspect a highly
chitinised conical projection; claspers bifid, both parts well
developed, pubescent on their basal halves and bearing also a few
longer hairs. Harpes: basal portions unequal, highly chitinised;
from the right basal portion, which is the broader, arises a rather
lightly chitinised posterior extension (shaped as shown in the figufe),
the distal end of which is twisted and covered by minute hairs.
Aedoeagus broadly-ending, the processes on each side of the
horizontal band highly chitinised, finger-like, and curved ventrally
at their tips.
NIGERIA (Northern provinces) : Kaduna, August, 1920 (Dr. W. B.
Johnson). This insect resembles D. -flava in some respects, but
is of an entirely different colour; the hypopygium of the male is
characteristic.
Dasyhelea boothi, sp. n.
Measurements.
Length of body (one male) . ro mm.
Length of wing .0*8 mm.
Greatest breadth of wing.0*25 mm.
Head dark brown. Eyes narrowly separated. Clypeus, proboscis
and palpi brown. Third palpal segment not inflated, about as long
as the fourth and fifth together, fifth about as long as the fourth.
Antennae : torus dark brown, flagellum brown; segments four to
eleven sub-spherical to ovoid, length from about once to once and
a third the greatest breadth; segments twelve to fifteen elongated,
sub-equal, but the fourteenth slightly the shortest, the twelfth,
thirteenth and fourteenth about three to four times as long as broad,
binodose, the fifteenth broader, not ending in a stylet. Thorax
dark brown. Scutellum pale brown, slightly darker at the sides;
bearing two lateral and four centro-marginal bristles, and one small
central hair. Post-scutellum dark brown. Pleurae brown. Wings
sparsely clothed with hairs which extend between the fourth and
fifth veins to the level of the cross-vein; venation as usual. Halteres
332
with brown stems and yellow knobs. Legs uniformly brown; claws
small, equal, simple, bifid at the tips. Abdomen dark brown.
Hypopygium (fig. 3, a to d). Ninth segment-, tergite sparsely
clothed with relatively short hairs, tapering distally, posterior
margin not notched and bearing double lateral processes; stemite
bare, prolonged posteriorly in the middle as a triangular process.
Forceps not very highly chitinised or hairy : side-pieces rather short
Fig. 3. a to d—Dasybelea bootbi , sp.n., outlines of male hypopygium ; ventral view :
a —ninth segment and forceps ; c —harpes ; d —aedocagus ; b —lateral view.
*—Dasybelea retorta , sp.n., spermatheca. (x 300 circa.)
and broad, with a hairy internal apical process; claspers moderately
chitinised, with somewhat squared ends, the proximal two-thirds
pubescent, bearing three short, stout, hairs about the middle of the
inner border. Harpes (fig. 3, b and c ) moderately well chitinised,
almost symmetrical, composed of a large and irregularly shaped
basal plate on each side, and a median common posterior extension
which is complex, expanded distally, and with a terminal ventral
333
barb-like process. Aedoeagus (fig. 3, b and d) very highly
chitinised in parts, apparently composed of a pointed median
process which is bent ventrally at its tip and in ventral view is
pyriform, two short and broad limbs, and external to them somewhat
T-shaped extensions.
NIGERIA (Cameroons): Victoria, April, 1921 (Dr. L. H. Booth).
We have pleasure in dedicating this species to the collector,
Dr. L. H. Booth.
Dasyhelea retort a , sp. n.
Measurements.
Length of body (one female) .1*35
Length of wing .0*9 mm,.
Greatest breadth of wing.0*3 mm.
Head dark brown. Eyes very narrowly separated dorsally.
Clypeus, proboscis and palpi dark brown. The third segment of
the palp cylindrical, not inflated, about as long as the fourth and
fifth segments together; the fifth segment shorter than the fourth,
widest at its distal end and rounded. Antennae dark brown, with
dark brown hairs and long curved spines on all the segments of the
flagellum; third segment slightly broader than the fourth; segments
four to fourteen forming a continuous series, gradually elongating
and becoming more flask-shaped, their length varying from about
once to once and a half the greatest breadth; the last segment
broader and longer, about three times as long as broad, ending in
a stylet. Thorax dark brown with small yellowish humeral patches.
Scutellum almost entirely yellow, sides only slightly infuscated;
bearing two lateral and four centro-marginal bristles, and a single
central hair. Post-scutellum dark brown. Pleurae brown, lighter
than the dorsum. Wings with decumbent hairs extending to the
base between the fourth and fifth veins. Costa reaching the middle
of the anterior border, and terminating beyond the bifurcation of
the fifth vein. First and third veins forming a small cell. Halteres
with yellow knobs with brown basal infuscation, and brown stems.
Legs light brown, with dark knee spots and slight infuscation of
the distal tarsal segments; hind femora slightly infuscated in the
middle dorsally. Claws simple, equal. Abdomen dark brown,
venter paler than the dorsum, yellowish pigmentation (which
334
disappears in caustic potash) visible laterally and between the
segments. Spermatheca single, highly chitinised, shaped something
like a chemical retort (fig. 3 e) ; distal portion sub-spherical,
length 44 a*, greatest breadth 38^; proximal portion, the chitinised
commencement of the duct, curved, arising obliquely, length about
30 /», width at its middle about up.
SlERRA Leone: Freetown, May, 1920; one female taken about
noon upon a window in the Royal Hotel.
Genus ATRICHOPOGON , Kieff.
Atrickopogon africanum t sp. nov.
Measurements.
Length of body (one female) .1*4 mm.
Length of wing -.1*3 mm.
Greatest breadth of wing.0*4 mm.
Head dark brown, with dark brown hairs. Eyes broadly
contiguous above, bare. Clypeus, proboscis and palpi dark brown.
First and fourth palpal segments short, second and third longer,
fifth somewhat dilated at its distal end; third segment slightly
inflated and furnished with a moderately large sensory pit in its
distal third. Antennae dark brown : segments four to ten short and
broad, the tenth sub-spherical; last five segments (eleven to fifteen)
elongate, sub-equal, four or five times as long as broad, the fifteenth
terminating in a relatively long stylet. Thorax uniformly dark
brown; clothed with a few short, dark-brown hairs. Pleurae dark
brown. Scutellum dark brown, but not so dark as the mesonotum,
bearing two sub-median and two lateral bristles and a few (about
half a dozen) short hairs. Post-scutellum dark brown. Wings
(fig. 4) generally similar to those of A. xanthoaspidium , unspotted
Fig. 4. Atricbopogon africanum , sp.n., outline of wing of female, (x 90 circa.)
335
but slightly infuscated, anteriorly somewhat darker than posteriorly;
anterior veins brownish. Surface uniformly covered with micro-
trichia, and bearing decumbent hairs, which, however, are more
scanty than in A. xanthoaspidium, none being present between the
branches of the fifth vein or in the anal fold, and only a very few
between the lower ramus of the fourth vein and the upper ramus of
the fifth. Petioiate portion of the fourth vein about the same length
as in A. xanthoaspidium. Halteres with dark brown knobs. Legs
almost uniformly brown, but distal segments slightly darker. First
tarsal segment of hind legs rather over three times as long as second.
Claws simple, equal, about half the length of the fifth tarsal segment.
Empodium well developed, as long as the claws. Abdomen dark
brown, ventral surface slightly paler. Spermatheca (fig. 5) single,
Fig. 5. Atricbopogon africanwm , »p.n., »permatheca. ( x 375 circa.)
large (length 107/1, greatest breadth 84/1), pear-shaped, heavily
chitinised; a short portion (about 15/1) of the duct, which is narrow,
is chitinised.
GOLD COAST: Accra, August, 1920, one female taken in the
evening upon a window in the laboratory.
Atrickopogon elektrophaeum , sp. n.
Measurements.
Length of body (one female) . ... 1*4 mm.
Length of wing .1*3 mm.
Greatest breadth of wing.0*5 mm.
Head yellowish-brown. Eyes contiguous above. Clypeus,
proboscis and palpi yellowish-brown. First segment of the palp
336
short; second, fourth and fifth sub-equal, about twice the length of
the first; third rather shorter than the combined lengths of the fourth
and fifth segments, slightly inflated in the middle, furnished with a
sensory cup opening at the junction of the middle and distal thirds;
fifth segment somewhat infuscated and tapering distally. Antennae
brown: first segment poorly chitinised, bearing several short hairs;
torus rather a light brown colour, sub-spherical, bearing a few short
hairs; third segment wider than any of the other basal segments;
segments four to ten short and broad, their lengths varying from
about three-quarters to a little over once the breadth; segments
eleven to fifteen elongate, cylindrical, sub-equal, four or five times
as long as broad, the last the longest and ending in a nipple-like
process. All the basal segments of the flagellum bear long, curved,
pointed spines. The ratio of the combined lengths of segments
three to ten to the combined lengths of segments eleven to fifteen as
o*6 to i. Thorax yellowish-brown, unicolorous, clothed with small
brown hairs and bearing larger and darker hairs above the wings.
Scutellum pale yellow-brown, bearing two sub-median and two
lateral bristles, and about half a dozen small hairs which are dark
brown in colour. Post-scutellum yellowish-brown, darker than the
scutellum. Pleurae yellowish-brown. Wings with a pale yellowish
tint, similar to those of A . xanthoaspidium but with fewer hairs,
none being present between the fourth and fifth veins nor between
the rami of the fifth vein, and only a row of eight along the false
vein in the anal angle. Halteres with pale yellow-brown knobs.
Legs almost uniformly yellowish-brown, terminal segments of the
tarsi, however, slightly darkened. First tarsal segment on all the
legs about three times as long as the second. Claws simple, equal,
about half the length of the fifth tarsal segment. Empodium well
developed, as long as the claws. Abdomen yellowish-brown, rather
darker than the thorax; venter paler than the dorsum. Spermatheca
single, highly chitinised, oval, large, length 106/1, greatest breadth
80/1 ; the duct not chitinised.
GOLD Coast : Accra, 1920; taken in the evening upon a window
in the laboratory.
337
Atrichopogon perfuscum , sp. nov.
Measurements.
Length of body .1*3 mm.
Length of wing .ri mm.
Greatest breadth of wing.0*4 mm.
Head dark brown, clothed with dark brown hairs. Eyes broadly
contiguous above, smooth. Clypeus, proboscis and palpi dark
brown. Antennae dark brown; segments four to ten short and
broad, the tenth sub-spherical the others broader than long, the
length varying from two-thirds to three-quarters the breadth; the
last five segments (eleven to fifteen) elongate, about three times as
long as broad. Thorax uniformly dark brown. Pleurae dark
brown. Scutellum dark brown, but rather lighter coloured than the
mesonotum, bearing two sub-median and two lateral bristles and
several (six to eight) short hairs. Post-scutellum dark brown.
Wings clear, unspotted; venation and decumbent hairs as in
A . xanthoaspidium , but fewer hairs present between the rami of the
fifth vein and in the anal angle (only two being present in the former
situation and one or two in the latter in one of the specimens
examined, rather more in the others). Halteres with buff-coloured
knobs and dark brown stems. Legs almost uniformly yellowish-
brown, but tarsal segments slightly infuscated. Claws equal, about
half the length of the fifth tarsal segment, each with a very small
sub-apical tooth. Empodium well developed, as long as the claws.
Abdomen dark brown. Spermatheca single, highly chitinised,
oval; length from 70 ft to 87/1, greatest breadth from about 57/4
to 65/4, the duct chitinised for only a short distance (about 5/4) at
its commencement.
Gold COAST: Accra, October, 1920; three females collected in
the evening upon the windows of the laboratory.
Atrichopogon chrysosphaerotum , sp. nov.
Measurements.
Length of body (two females) .1*15 mm.
Length of wing .0*9 mm.
Greatest breadth of wing.0*35 mm.
Head dark brown, clothed with dark brown hairs. Eyes broadly
contiguous above, bare. Clypeus, palpi and proboscis dark brown.
338
First palpal segment short; second, third and fourth longer, sub¬
equal; fifth rather shorter; third segment moderately inflated
and furnished with a well developed sensory pit. Antennae dark
brown, segments four to ten short and broad, the tenth sub-spherical,
the others broader than long, the length varying from half to about
two-thirds the breadth; last five segments (eleven to fifteen)
elongate, from two and a half to three times as long as broad, the
fifteenth terminating in a relatively large stylet. Thorax uniformly
dark brown, scantily clothed with brown hairs. Pleurae dark
brown. Scutellum dark brown, bearing two sub-median and two
lateral bristles and no short hairs. Post-scutellum dark brown.
Wings clear, unspotted, the anterior veins brownish; venation as in
A. xanthoaspidium , but first cell larger; decumbent hairs absent,
wing surface covered by microtrichia. Halteres with yellow knobs
and pale straw-coloured stents. Legs almost uniformly yellowish-
brown, but tarsal segments slightly infuscated. Fourth tarsal
segments not cordiform; first tarsal segment of hind legs nearly four
times as long as second; dorsal hairs on the tarsal segments rather
long. Claws equal, apparently simple or with a very minute sub-
apical tooth, about half the length of the fifth tarsal segment.
Empodium well developed, as long as the claws. Abdomen dark
brown with a yellowish tint, pigmented with a substance which does
not clear in carbolic acid but which is removed by caustic potash.
Spermatheca single, highly chitinised, oval; length 57 p, greatest
breadth 46/1, only the very commencement (about 4 fi) of the duct is
chitinised.
Gold COAST: Accra, November, 1920; a single female, collected
in the evening upon a window in the laboratory. Oblogo, May,
1920, reared from rotten wood from a canoe in the river Densu.
Atrichopogon homoium , sp. nov.
This species, of which we possess only a single female, agrees
with the foregoing (A. chrysosphaerotum, sp. nov.) in size, coloura¬
tion, and apparently in every other particular excepting in the
distribution of the hairs on the wings and the scutellum. On the
scutellum are four short hairs, two on each side, in addition to the
bristles. On the wings are a few decumbent hairs; seven to ten at
339
the tip of the wing, one near the periphery between the rami of the
fourth vein, and a few along the distal portion of the anterior ramus
of the fourth vein (fig. 6).
Fig. 6 . Atricbopogon bomoium , sp.n., wing of female. (x 120 circa.)
The distribution of the hairs on the wings and the scutellum
appear to be important specific characters, and therefore, notwith¬
standing the similarity of this insect to A. chrysosphaerotum and
the paucity of our material, we feel compelled to regard it as
a separate species.
GOLD Coast*. Oblogo, September, 1920; a single female reared
from material taken from a canoe.
Genus KEMP 1 A , Kieff.
This genus, originally described by Kieffer (1913) as a sub-genus
of Dasyhelea and subsequently raised to generic rank by its author
and removed to the Atrichopogon group, appears to be characterised
chiefly by the presence of a well developed empodium on the legs
and of pubescence on the eyes, and the absence of the longer hairs
from the wings. We have referred the insect described below to the
genus Kempia , notwithstanding the fact that it bears on its wings
a few decumbent hairs, because, as stated elsewhere, we are inclined
to regard this character as of specific rather than generic value.
It may be noted, however, that in the case of Prokempta the absence
of the longer wing hairs appears to have been considered by Kieffer
sufficient justification for its separation from Dasyhelea.
3+0
The larvae and pupae resemble those of F orcipomyia. The
larvae were found living upon the water lettuce Pistia stratiotes,
hut were normally seldom seen alive, probably because they were
buried in the substance of the plant. The pupae, the posterior
portions of which are enveloped by the larval pelts, were found
partially embedded in the leaves and between the papillae which
cover densely the surface of the leaves. They appeared to be
sedentery.
The genus, therefore, shows affinities to three different types
of the Ceratopogortinae , the characters of the wing linking it to
Atnchopogon, those of the eyes to Dasyhelea, and those of the early
stages to F orcipomyia.
Kempia ochrosoma, sp. nov.
Measurements.
Length of body . . I '$ mm.
Length of wing . . ... ... n nun.
Greatest breadth of wing ... ... ... ... . 0’35 mm.
Head straw-yellow, clothed with yellow hairs. Eyes pubescent:
in the female contiguous, but with the facets narrowly separated;
in the male separated. Clypeus, palpi and proboscis pale straw-
yellow, with similarly coloured hairs. First palpal segment small,
second and fourth sub-equal, fifth short and somewhat expanded at
its distal end, third longer than any of the others, slightly inflated
in its middle third, and furnished with a deep sensory cup.
Antennae : first segment and torus straw-yellow; in the female,
flagellum segments slightly infuscated, especially the more distal
ones, segments eleven to fifteen dark brown; in the male, segments
three to eleven pale yellow, segment twelve dark distally, and the
last three segments dark brown; whorls of hairs pale straw-yellow.
In the female, segments four to ten sub-spherical, the length varying
from a little less than to about the same as the breadth, segments
eleven to fifteen elongate, from twice to three times as long as broad,
the last segment terminating in a stylet. In the male, the last three
segments elongate, sub-equal, the thirteenth slightly the longest and
the fifteenth terminating in a stylet. Thorax uniformly coloured,
ochraceous, almost the same hue as the head; clothed with short,
curved, yellow hairs dorsally, and with a few longer, spine-like.
“V
34 1
yellow hairs posteriorly. Pleurae ochraceous. Scutellum slightly
paler yellow than the mesonotum, bearing two admedian and two
lateral bristles and a few (ten in the female, eight in the male) short
hairs. Post-scutellum ochraceous. Wings unspotted, the anterior
veins yellowish; venation as shown in the figure (fig. 7). Decumbent
hairs scanty: in the female, limited to the distal third of the wing
and a few near the posterior margin; in the male, almost entirely
wanting. Surface of the wing closely covered with microtrichia.
Halteres with pale straw-yellow stems and white knobs. Legs
almost uniformly pale straw-yellow; unarmed. Claws in both sexes
equal, about half the length of the fifth tarsal segment; in the female,
each with a very small, sub-apical tooth, in the male, deeply
bifurcated at the tips. Empodium well developed, hairy; at least
as long as the claws. Add omen ochraceous, rather paler yellow than
the thorax. Hypopygium of the male of a similar colour and feebly
chitinised. Spermatheca single, rather feebly chitinised, obovate
and very large; length about 230 ft, greatest breadth about 150/*,
the duct chitinised at its commencement for a short distance
(about 10ft). The abdomen, and also the thorax and the knobs of
the halteres, containing a yellowish pigment which is not cleared by
carbolic acid but which dissolves in caustic potash.
Hypopygium (fig. 8 a and b ). Ninth segment feebly chitinised;
tergite rather short and hairy, posterior margin rounded, without
lateral, finger-like processes. Forceps : side-pieces normal, feebly
chitinised; claspers also feebly chitinised, about as long as the
34 2
side-pieces, tapering distally, hairy almost to their tips. Harpes
apparently absent. Aedoeagus feebly chitinised, a large ventral
median structure with a central and two lateral processes; -when fully
expanded as shown in fig 8 a t but perhaps more frequently appearing
as in fig. 8 b. We are not able at present to suggest what are the
homologies of the median structures.
PUPA. Length about 2*1 mm., feebly chitinised; the posterior
half of the abdomen (from the fifth segment) enclosed within the
larval pelt, which is much shrivelled and discoloured. Integument
irregularly covered with rather sparsely scattered spicules. Cephalo -
thorax relatively large and broad, somewhat of the Forcipomyia
type but with no dorsal extension over the middle of the first
abdominal segment. Respiratory trumpets almost smooth, short
and nearly straight, length a little less than 0*2 mm.; they arise from
343
small tubercles and are without stalks. Main tracheal trunk does
not give off any lateral branches, and ends distally in a double row
of rather long, blunt processes. Operculum feebly chitinised,
sparsely spiculated; at its posterior angle is a small tubercle covered
with long spicules. Anterior marginal tubercle an irregularly
conical elevation covered by long spicules, and bearing a long
bristle; anterior dorsal situated posterior and external to the anterior
marginal and anterior to the base of the trumpet, a small tubercle
bearing a rather long hair; dorso-lateral apparently unarmed,
except by spicules, ventro-lateral apparently absent or obsolete;
ventro-median very small, bearing a short hair. Dorsal tubercles
remarkable, six on each side, each giving rise to a delicate,
branching, dendritic process resembling a sort of gill (fig. 8 d );
these processes are well developed on all the tubercles excepting the
small admedian pair situated a tittle anterior to the most posterior
pair; two of these tubercles are armed, the two situated most
anteriorly and internally, and bear long spines. Abdomen directed
straight backwards, tapering rather rapidly, the last segment
terminating on each side in an almost straight process which is
slightly dilated at its distal end, and is covered with long spicules
directed anteriorly, which no doubt function as hold-fasts (fig. 8 c ).
The segments, which are enclosed within the larval pelt, bear only
rudimentary tubercles; on the other segments (one to four) are a pair
of dorsal admedian, and a pair of lateral tubercles similar to those
on the dorsum of the cephalo-thorax—that is, dendritic processes
resembling gills—the dorsal ones bear large spines.
LARVA. Length about 3 mm. when fully grown, brownish in
colour. Head moderately chitinised, more or less conical, bearing a
few relatively large hairs or bristles. Hom-like appendages on the
dorsum (which are connected with the distal ends of the
antennae of the adult) small, narrow, about 0 06 mm. long, straight,
arising from quite small tubercles. Eyes large, bilobed. Mandibles
densely chitinised, black, terminating in three or four teeth.
Hypopharyngeal sclerite very highly chitinised, the posterior part
armed on each side with a comb-like row of about ten short, pointed
teeth. Body composed of twelve visible segments; cuticle covered
with coarse spicules. Armature of bristles, spines, etc., modified at
the anterior and posterior ends, but general arrangement as follows.
344
A pair of finger-like, admedian dorsal tubercles, bearing long spines,
which are more or less barbed at their bases; three pairs of very long
(about o*8 mm.) and delicate dorso-lateral processes; four pairs of
ventral spines, two central, the one admedian and the other ventro¬
lateral, and two postero-marginal, both ventro-lateral, these spines
are relatively short and are freely barbed. The long dorso-lateral
processes are the most conspicuous features of the larva, they bear
lateral spines or bristles, are directed backwards, and trail behind
the larva. The pro-thoracic pseiidopods are partially fused, highly
spiculated, each armed with a group of about ten large, well
developed hooks, and anteriorly with numerous small hooks. The
anal pseudopods are armed with similar large hooks arranged in two
transverse rows.
Gold COAST: Oblogo, February to March, 1921; reared from
plants of the water lettuce (Pistia stratiotes) taken from a swampy
pool, and from backwaters of the river Densu (PI. XXI, fig. 1).
Genus MONOHELEA , Kieffer.
This genus is stated by its author to possess the characters of
Stilobezzia , Kieff., but the petiole of the fourth longitudinal vein is
very short, the fourth tarsal segment is long and cylindrical, the
claws of the fore and middle legs of the female are simple, equal,
two-thirds the length of the last tarsal segment, and the claws of the
hind legs single, longer than the last tarsal segment. The species
described below, of which at present we possess only a single
female, appears to belong to this genus, although not conforming
exactly to the generic description given above (for example, the
claws of the fore and middle legs are not simple). The insect
superficially resembles Stilobezzia , and when at rest holds its wings
in a manner similar to 5 . spirogyrae , that is, diverging slightly, and
not folded one on top of the other on the dorsum of the abdomen.
Monohelea litoraurea , sp. n.
Measurements.
Length of body (one female) . i # 3 mm.
Length of wing .ronun,
Greatest breadth of wing.0*4 mm.
Head : occiput dark grey to brown, clothed with dark brown
hairs. Eyes smooth, contiguous dorsally. Clypeus and proboscis
345
dark brown. Palpi dark brown, third segment slightly inflated and
furnished with a sensory cup; fifth segment rather long and narrow,
with a terminal group of four hairs. Antennae darkish brown: first
segment rather large, dark brown, bearing a few dark hairs; torus
dark brown, bearing a few dark hairs; flagellum segments paler,
gradually elongating and deepening in colour towards the distal end
of the antenna; fourth to tenth segments sub-cylindrical, from about
one and a third times to twice as long as broad; segments eleven to
fifteen elongate, from about three to a little over four times as long
as broad, the last segment being the largest and the longest and
tapering at its extremity, but not terminating in a definite stylet.
Whorls of hairs small and the constituent hairs short; there is a hair
just before the tip of the last segment. Thorax : dorsum dark grey
with many dark brown spots and patches, and with the antero-
Fig. 9. Monobelea litoraurta , sp.n., wing of female. ( x 105 circa.)
lateral angles brownish-yellow. The arrangement of the spots and
patches is somewhat like that on the thorax of Culicoides schultzei .
The mesenotum is clothed with rather short, dark hairs. Pleurae
dark brown. Scutellum darkish brown, but not so dark as the
mesonotum, paler in the middle than at the sides, bearing two
admedian and two lateral bristles and a few (six) short hairs. Post-
scutellum dark brown. Wings grey, with large white patches; the
arrangement of the patches and the venation as shown in fig. 9.
Lower ramus of fourth vein obsolete at the proximal end. The
surface of the wing is covered by microtrichia, but without
larger decumbent hairs. Halteres pale, with white knobs. Legs
brown, femora and tibiae more or less infuscated. Fore and middle
34 6
femora brown, somewhat infuscated, especially at their apices, not
swollen and without strong spines; hind femora uniformly very dark
brown, slightly swollen, without strong spines. Fore and middle
tibiae brown, infuscated; hind tibiae uniformly very dark brown,
somewhat swollen. Tarsal segments brown, paler than the proximal
segments: fore and middle tarsi without regularly arranged rows of
spines, fore tarsi with a well developed black spine at the apex, the
base, and sometimes the middle of the first segment and somewhat
smaller but similar spines at the apex of the second and third
segments, middle tarsi with several similar but less well defined
spines, on the first segment and at the apex of the second and third
segments, hind tarsi (fig. ioa) with a regularly arranged ventral
row of small spines on the first segment, and with large black spines,
one at the apex and one at the base of the first segment, two at the
apex of the second segment, one at the apex of the third segment,
and t>vo at the apex of the fourth segment. Fourth tarsal segment
on all the legs cylindrical. Claws on the fore and middle legs small,
about half the length of the fifth tarsal segment, equal, each with a
small basal tooth and a bifid extremity; claws on the hind legs
single, with a very long tooth (as long as the fourth and fifth tarsal
segments together), and a short tooth apparently fused with the
large tooth at the base. Empodium rudimentary. Abdomen very
dark brown, excepting at its extreme proximal end, where it is
yellowish-brown; distal end almost black. Dorsal surface very
sparsely clothed with short, dark hairs, which are almost entirely
restricted to the sides. Spermathecae (fig. io b ) two, unequal,
347
highly chitinised, pyriform (measurements about 8 op by S 5 M, and
65 fi by 55/4); the commencement of the duct, which is narrow
(about 4/1), is chitinised for some distance (about 20/*).
Gold Coast: Accra, 27th March, 1921; collected on a window
in the laboratory at 6 p.m.
Genus EUKRAIOHELEA , nov.
The two following species show kinship to the genus Stilobezzia ,
Kieff., the chief generic characters of which are, according to
Kieffer (1919)—wings glabrous, the first and third veins forming two
radial cells of which the second is the longer, the fourth vein
petiolate, the fourth tarsal segment cordiform in both sexes, and the
claws long, simple, and very unequal in the female, short and equal
in the male. Thy possess the above characters excepting that the
first radial cell of the wing is obsolete, the first and. third veins
forming only a single large cell, but they show certain other
divergencies from earlier descriptions given by Kieffer (1917), for
example, the fore femora are armed but not swollen, and the hind
tibiae beai spines. Moreover, the hypopygium of the males, whilst
closely resembling one another, diverge from the type found in
Stilobezzia spirogyrae. These morphological differences are, in our
opinion, of such a nature as to necessitate the separation of these two
species from the genus Stilobezzia , and, therefore, notwithstanding
their close similarity in other respects to species of that genus, we
propose to regard them as belonging to a new genus, for which
we suggest the name Eukraiohelea .
Measurements.
Eukraiohelea africana , sp. n.
Male.
Female.
Length of body
Length of wing
Greatest breadth of wing
i # 3 mm. 1*4 mm.
1*2 mm. 1*3 mm.
0*4 mm. o*s mm.
The ground colour of this insect is olive green, on which are
superimposed the brown markings. Head brown. Eyes smooth;
narrowly separated in the female, more widely in the male. Clypeus
and proboscis dark brown. Palpi dark brown, moderately long,
the third segment slightly inflated distally and furnished with a
shallow sensory cup near its apex. Mouth-parts well developed in
the female. Antennae : first segment small; torus yellowish-brown,
sub-globular, very large in the male, bearing a few short hairs;
flagellum pale brown proximally, the last three segments in the male
and five in the'female dark brown. Plume of the male moderately
well developed, pale brown; hairs of the flagellum of the female very
short, brown. In the male, the last three segments of the flagellum
elongated, the thirteenth and fourteenth about five and twelve times
as long as broad respectively, the fifteenth much longer, about
thirty times as long as broad, and ending in a short, stout process.
In the female, segments four to ten pale brown proximally, dark
brown distally, and segments eleven to fifteen all dark brown;
segments four to ten sub-cylindrical, rather wider in the middle
than at either end, sub-equal, about three times as long as broad;
segments eleven to fourteen elongated, about ten times as long as
broad; the last segment much longer, about twenty times as long
as broad, ending in a short, stout process. Thorax blue-green, with
brown infuscation on the dorsum; hairs large, scanty, dark brown.
Pleurae blue-green \ a dark brown spot on the coxae of the fore and
middle legs. Thoracic pits absent. Scutellum blue-green in the
middle, and brown, with a blue-green tint, at the sides; bearing two
admedian and two lateral bristles and a few (two to four, but often
absent in the male) short hairs. Post-scutellum dark brown with a
greenish-blue tint. Wings unspotted, without decumbent hairs.
Wing surface covered by microtrichia. Venation as shown in the
figure (fig. ii); first radial cell obsolete; in the female, the fourth
349
vein forks distally to the fifth, in the male, they both fork at about
the same level. Fringe very short on the distal part of the wing.
Halteres with greenish-blue knobs; stalks almost white, bases of the
knobs infustated. Legs pale brown, almost colourless, with dark
knee spots; the infuscation on the hind legs extending beyond the
knee nearly half way down the tibia. Fore femora armed with two
short, stout, ventral spines, middle and hind femora unarmed; hind
femora slightly swollen, fore and middle normal. Fore tibiae with
the usual long apical spine; hind tibiae with a ventral row of spines,
three long ones on the middle .third and four short ones more
distally. On all the legs the first tarsal segment at least twice as
long as the second, third small and almost cordiform, fourth very
small and strongly cordiform, fifth slightly infuscated, about as long
as the second and longer than the third and fourth together. Rows
of bulbous spines present on the first and second tarsal segments of
all the legs; a stout basal spine on the first tarsal segment of the
middle and hind legs; on the proximal half of the fifth tarsal
segment of the fore and middle legs are two pairs of stout spines,
on the hind legs a single pair. Claws alike on all the legs: in the
female, single, with a long tooth (as long as the fifth tarsal segment),
and a tooth about half this length apparently fused with it at the
base, which is, moreover, somewhat extended as a process; in the
male, shorter (about two-thirds the length of the fifth tarsal
segment), bifid at the tips, composed' of two equal parts fused
at their bases. Empodium absent. Abdomen green,with brown
infuscation (which is most marked in the female) at the sides
and posterior dorsal margins of the segments. First segment
with conspicuous lateral tufts of hairs. Spermathecae two, highly
cHitinised, pyriform (about 46 ft by 35 a 1 ); the commencement of the
duct chitinised for a short distance (about 7 n) only.
Hypopygium (fig. 12). Ninth segment : tergite rather feebly
chitinised, moderately hairy; posterior margin broad, not cleft,
without finger-like lateral processes; apical lobe-like processes
moderately well developed, hairy; sternite apparently not excavated
in the middle posteriorly. Forceps : side-piece's moderately long
and hairy, distal ends infuscated; claspers rather broad, not strongly
chitinised, tips infuscated, clothed with very delicate minute hairs
and bearing a few longer hairs. Harpes in form somewhat
35 °
resembling those of species of Dasyhelea, but with the distal
extensions bilateral and separate. Basal portions irregularly
shaped, broad laterally; posterior extensions long, lath-like
processes directed almost straight backwards, with bluntly pointed
tips, reaching posteriorly as far as, or a little further than the margin
Fig. 12. Eukraiobeha africana , sp.n., outline of male hypopygium, ventral view.
( x 375 circa.)
of the ninth tergite. Aedoeagus apparently composed of two
separate chitinised lateral rods corresponding to the limbs of the
arch in other genera, and a delicate membranous portion enclosing
them and prolonged posteriorly in the middle line as a conical
process. Ventral wall not spiculated.
GOLD Coast : Swamp between Koforidua and Tafo, a little north
of Accra, April, 1921; Weshiang, June, 1921; reared from plants of
the water-weed Pistia stratiotes .
35 1
Eukraiohelea versicolor , sp. n.
Measurements.
Male.
Length of body
Length of wing
Greatest breadth of wing
1*3 mm.
l*o mm.
o*3 mm.
Female.
1*3 mm.
i*i mm.
o *4 mm.
This insect resembles the last species, but the ground colour is
white instead of blue-green. In the following account, only the more
important points of difference between it and Eukraiohelea africana
will be given.
Head dark brown; occiput dark brown in the middle,
paler at the periphery; median occipital hairs long, dark brown.
Eyes during life metallic green; smooth, separated in both sexes.
Proboscis dark brown, well developed in the female. Palpi dark
brown, as in E. africana. Antennae : torus yellowish-brown;
flagellum very pale brown at the base, last five segments in the
female and three in the male completely dark brown, apical portions
of the fourth to the tenth segments in the female infuscated. Plume
in the male moderately well developed, pale brown, hairs in the
female short, pale brown. In the male, the last three segments
elongated, about five, twelve, and twenty-two times as long as
broad respectively, the last segment tapering to a conical tip. In
the female, segments four to ten as in E. africana ; segments eleven
to fourteen elongate, sub-equal, about eight or nine times as long as
broad; segment fifteen longer, about twelve times as long as broad,
ending in a conical tip. Thorax : ground colour white; anterior
half of the dorsum almost entirely dark brown in the male,
in the female more or less infuscated and dark brown, but
with median pale areas on each side of the middle line; in
front of the scutellum is a triangular dark brown mark with its base
directed posteriorly. Pleurae white. Over the bases of the coxae is
a small, oval, dark brown patch. Scutellum dark brown, armature
of bristles and hairs as in E. africana Post-scutellum dark brown.
Wings as in E. africana. Halteres white. Legs : ground colour
white; distal half of the hind femora and apical sixth of the
hind tibiae dark brown; fifth tarsal segments hardly infuscated.
Coxae infuscated. Femora shaped and armed as in E. africana u
Tibiae shaped and armed as in E. africana , but hind tibiae bearing
only three long spines. Tarsal segments as in E. africana ; spines
352
on the fifth segments, however, less well developed, arrangement in
the female as in E. africana , in the male only one pair present on
all the legs. Claws as in E. africana . Abdomen white, with dark
brown markings arranged as follows: small lateral patches on the
first, fourth, and seventh segments, large dorso-lateral patches,
reaching almost to the middle line dorsally, on the second, third,
fifth and sixth segments. Lateral hair tufts on the first segment
not so prominent as in E. africana. Spermathecae similar to those
of E. africana ; two, highly chitinised, pyriform (about 35 p by 30/4),
the commencement of the duct chitinised for a short distance
(about 4 p) only.
Hypopygium (fig. 13). Hypopygium darkish brown, rather
small. Ninth segment : tergite moderately hairy, tapering slightly
distally, posterior margin nearly straight, with a trace of a median
Fig. 13. Eukraiobelea versicolor 9 sp.n., outline of male hypopygium, ventral view.
(x 375 circa.)
cleft, but without lateral finger-like processes; apical lobe-like
processes well developed, hairy; sternite apparently prolonged
posteriorly as a delicate, median, cone T shaped process. Forceps :
side-pieces rather short and stout, moderately hairy; claspers long,
stout, feebly chitinised, especially at their distal ends, entirely
353
covered by very delicate minute hairs and bearing at the posterior
extremity a few rather larger hairs* Harpes similar to those of
E. africana , but basal portions more expanded laterally, and
posterior extensions longer and bent sharply in an anterior direction
at about their middles. Aedoeagus somewhat similar to that of
E. africana , delicate membranous part apparently with large lateral
folds; ventral wall not spiculated.
GOLD COAST: Swamp between Koforidua and Tafo, a little
north of Accra, April, J921; reared from plants of the water-weed
Pistia stratiotes .
Genus SCHIZODACTYLUS, nov.
This genus is allied to Xylocrypta , Kieff., and Xenohelea , Kieff.,
genera which have been separated by Kieffer (1917) from
Sphaeroptias (Stephens), Curtis, and Palpomyia t Mergele, by the
characters of the fourth tarsal segments, which are cylindrical in
both sexes, and the antennae of the males, only the last three
segments of which are elongated. From the former it may be
distinguished by the facts that the eyes in the male are separated
and the body is not squat; from the latter by the fact that the claws
of the female are equal. The chief generic characters are as
follows:—Eyes smooth, separated in both sexes; widely in the male,
narrowly in the female; last three segments of the antenna of the
male elongate; wings covered by microtrichia but without longer
decumbent hairs, costa reaching beyond the middle of the wing,
first and third veins forming two radial cells, the distal of which is
the longer, cross-vein not very oblique and not twice as long as the
base of the cubitus, fourth vein sessile; femora armed, fourth tarsal
segments cylindrical, claws in the female long and equal, those on
the fore legs with long basal barbs, empodium rudimentary.
Schizodaetylus telmatoscopus } sp. n.
Measurements.
Length of body (two males) .2*8 mm.
Length of wing .1*5 mm.
Greatest breadth of wing.0*5 mm.
Head dark brown, large, wider than the thorax. Eyes glabrous,
widely separated. Proboscis dark brown, very short. Palpi
554
brown, very small; third segment not inflated, sensory pit small or
rudimentary. Antennae rather dark brown, especially the torus and
the three terminal segments. First segment without hairs; torus
sub-spherical, very dark; third segment rather larger than the
followjng segments; fourth to twelfth segments almost cylindrical,
from one and a third to two and a half times as long as broad, sharply
separated from one another; last three segments elongated, the
fifteenth being the longest and not ending in a stylet. Hairs not
very long, pale coloured, arranged somewhat irregularly and not
forming a single whorl. Thorax uniformly dark brown; small
pro-thoracic lobes present; no tubercle on the front margin of the
thorax in the middle; dorsum almost devoid of bristles, and without
either anterior or posterior pit-like depressions. Scutellum dark
brown, bearing a few (twelve to fourteen) hairs, but no large
bristles. Post-scutellum dark brown. Pleurae dark brown. Wings
unspotted and without long decumbent hairs; surface closely
covered with minute upright spicules; fringe short; stronger Hairs
on costa scanty. Venation as shown in the figure (fig. 14); first cell
Fig. 14. SfbizoJaciylut ttlmatttcopus, tp.n., venation of wing of male. ( x 70 circa.)
rather large and long. Halteres with pale brown knobs. Legs-.
femora and tibiae dark brown; first four tarsal segments pale
coloured with slightly infuscated apices, last segment entirely dark.
Trochanters with small, paired, curved spines! Femora not
unusually broad, bearing on all the legs two or three stout, short,
dark spines on the under surface near the apex. First tarsal
segment about twice the length of the second on all the legs; last
segment rather elongated, fourth not cordiform. Regular rows of
small spines are situated ventrally on the first tarsal segments of
355
the middle legs and on the first and second tarsal segments of the
hind legs; those on the first tarsal segments of the hind legs arranged
in a double row. Apically the fore tibiae bear a long; pale-coloured,
ventral spine, the middle tibiae a strong, dark-coloured spine, and
the hind tibiae the usual double row of bristles. First four tarsal
segments of the middle and hind legs with a pair of strong, dark
spines apically, those on the fourth segments being more slender.
Claws about half the length of the fifth tarsal segment, equal,
simple, but with bifid ends; empodium rudimentary. Abdomen
dark brown; venter slightly paler than the dorsum.
Hypopycium (fig. 15). Dark brown, well chitinised, relatively
rather small. Ninth segment : tergite bearing few bristles,
terminating distally on each side in a large hairy process bearing
two relatively long bristles; sternite deeply excavated in the middle,
Fig. 15. Scbizodactylus telmatoscopus , sp.n., male hypopygium. a—donal view ;
b, c and d y ventral view*, a —ninth tergite (small hair* not shown); b —harpes;
c —aedoeagus; d —forceps (hair on side piece not shown), (x 150 circa.)
reduced to a narrow band of chitin. Forceps : side-pieces rather
narrow, hairs not very long; claspers short, bearing at the distal end
a small claw. Harpes fused into a median, strongly chitinised rod;
proximal extremity bifurcated, each half with two processes; distal
end expanded, rounded in ventral view but slightly spoon-shaped in
lateral view, not highly chitinised, closely covered with minute hairs.
Aedoeagus : Y-shaped, chitinised portion with lateral flaps of
spiculated membrane on each side; distal end broad, with a small
35 $
spine at each side, and an irregularly chitinised fringe extending
anteriorly; ventral wall between the limbs of the Y well chitinised,
speculated; membrane connecting the aedoeagus with the ninth
sternite with a very few spicules at its distal border only.
FEMALE. The following morphological details of the female
were made out in a specimen extracted from a pupa of the same
sample, and which was apparently identical with those of the males.
Head : eyes narrowly separated. Palpi longer than in the male,
and segments relatively more slender; first segment small, second,
third, and fifth longer and sub-equal, fourth somewhat shorter.
Third palpal segment not inflated, without a definite sensory* cup
but with a slight anterior depression from which arise a number of
sensory hairs, the extremities of which are only slightly dilated.
Antennae : first segment bearing four hairs; torus somewhat pyriform
and bearing a considerable number of short hairs; segments four to
ten cylindrical, from about twice to three times as long as broad;
segments eleven to fifteen elongate, sub-equal, about six times as long
as broad; last segment not terminating in a stylet. Thorax : scutellum
bearing numerous (about thirty-three) long, bristles of somewhat
unequal sizes and arranged more or less in three rows in thk middle
and in two rows laterally. Legs : fore femora with ventral row of
twelve to fourteen short, stout, dark spines on its apical half; middle
and hind femora with about a dozen similar spines, some of those
situated nearest the apex being paired. Fore, tibiae with a long,
feebly chitinised, ventral spine apically; middle tibiae with short,
dark, strongly chitinised spine in a similar position. Fifth tarsal
segment of all legs with two pairs of stout, dark spines at the base;
middle legs, and hind legs less distinctly, with a pair of somewhat
similar spines at the apex of the first, second, and third tarsal
segments. Double row of small spines on the first tarsal segment of
the hind legs complete. Claws long, equal, almost as long as the
fifth tarsal segment; those of the fore legs (fig. 16) with a rather
large basal barb, which is not present on the other legs. Empodium
rudimentary. Abdomen : spermathecae two, highly chitinised,
oval; length 123/1 to 133/“, greatest breadth 106/1 to 121/1; only the
very commencement (about 4/1) of the duct chitinised.
PUPA. Length about 4 mm. to 5 mm. Form similar to that of
Culuoides t and, therefore, the description will be given on the same
357
lines as was done in the case of that Genus. The pupa is very dark
coloured and highly chitinised, especially at the anterior part of the
cephalo-thorax. Respiratory trumpets short, broad, and straight,
arising from rather depressed tubercles and without definite stalk;
length about 260/1, breadth in the middle about 65/1. The main
tracheal trunk is very broad, straight, without lateral branches, at
its distal end terminating in a semi-circular, fan-like arrangement of
about fifteen short processes. Cephalo-thorax . Anterior marginal
tubercle small, bearing a small bristle; anterior dorsad small,
bearing a small bristle; dorso-lateral small, bearing two bristles;
ventro-lateral small, bearing two or three short bristles; ventro-
median obsolete, represented by two small hairs. Dorsal tubercles
Fig. 16. Scbizodactylus telmatoscopus , sp.n., fifth tarsal segment and claws of
foreleg of female.
practically obsolete: anterior represented by two bristles slightly
separated from one another; posterior by a single bristle; and lateral
by a bristle and a socket-like mark, apparently unarmed, a little
external to it. The dorsum is highly chitinised and very dark, the
integument closely covered by small dark granulations; there is a
transverse band of dark patches on the anterior third similar to that
seen on many species of the Genus Dasyhelea. The postero-dorsal
tubercle apparently obsolete. The posterior margin of the dorsum
is rounded, not prolonged backwards as a median process.
Abdomen : first segment, very short, second, long and broad; third
to eighth, sub-equal but tapering towards the posterior extremity,
358
almost square; ninth, short and small, with two short* sharply-
pointed, divergent terminal processes. There are a few dark spots
on the integument of most of the segments : dorsally, two sub-lateral
anterior, two admedian central, and a central a little farther back;
ventrally, two sub-lateral anterior, and a central somewhat heart-
shaped spot between and behind them. The tubercles are all small*
and bear only small spines or hairs. The following may be
distinguished on each side of a typical segment. Dorsal tubercles:
antero-submarginal, two, very small, the outer being situated almost
laterally; postero-marginal, three, small, the inner one situated
posterior to the inner antero-submarginal tubercle, the outer two
contiguous, situated almost laterally. Lateral tubercles: antero-
submarginal, single, very small; postero-marginal, two, rather larger
than the other tubercles, situated close together and rather more
anteriorly than the corresponding dorsal and ventral tubercles, the
dorsal one appears to be composed of two fused tubercles and bears
a small spine and a short hair. Ventral tubercles: three, small,
postero-marginal in position and situated almost laterally; the inner
two are the larger and are contiguous, the outer one is very small
and is situated slightly more anteriorly than the others.
Gold Coast: Accra, December, 1920; numerous pupae (of
which only two, males, hatched), found in puddles of dirty water
near a stand-pipe beside the Weshiang railway line, about two and
a half miles from Accra.
Genus SPHAEROMIAS (Stephens), Curtis.
This genus includes those midges in which the eyes are
bare, the wings bare except for minute, point-like hairs visible
only with a microscope, third vein extending beyond the middle
of the wing, second radial cell longer than the first, fourth vein
forking almost under the cross-vein, femora unarmed and not
swollen, fourth tarsal segments cordiform, fifth not swollen,
empodium absent or rudimentary. It is apparently the same as the
Genus Johannsenomyia erected by Malloch to include those species,
which he had previously included in Johannseniella, * which have the
media furcate proximad to the cross-vein.’ One male of a single
species of this genus was obtained near Accra.
359
Sphaeromias litoranrea , sp. nov.
Measurements.
Length of body (one male) . 1*6 mm.
Length of wing . . V2 mm.
Greatest breadth of wing.0*4 mm.
Head dark brown. Eyes widely separated, smooth. Clypeus
and proboscis dark brown. Palpi dark brown: all five palpal
segments short, but the third and the fifth rather longer than the
fourth; third palpal segment slightly longer than broad, not inflated,
with a few long, knob-ended sensory hairs on its inner side anteriorly
which arise from a very shallow depression. Antennae dark brown :
plume poorly developed, composed of relatively few and short hairs
which are not arranged in distinct whorls; segments four to eleven
progressively longer, from once and a half to a little over twice as
long as broad; segment twelve about three times as long as broad;
segments thirteen to fifteen longer, sub-equal, bearing only short
hairs, the terminal segment tapering distally but not ending in a
stylet. Thorax uniformly dark brown. Pleurae dark brown.
Scutellum dark brown, bearing two admedian, two sub-median, and
two lateral bristles. Post-scutellum dark brown. Wings clear,
unspotted, the anterior veins brownish. Third vein extending some
distance beyond the middle of the wing; two radial cells, both well
formed, the first rather large, rectangular, the second longer than the
first; fourth.vein bifurcated, sessile, the bifurcation taking place a
little proximal to the cross-vein. Wing surface covered by micro-
trichia but otherwise bare, without decumbent hairs. Halteres
with dark brown knobs. Legs almost uniformly brown, but with
indications of darker knee-spots and with the tarsal segments slightly
infuscated. Trochanters with usual pair of stout, curved spines.
Femora unarmed, not swollen. First tarsal segments much longer
than second, fourth cordiform. First and second tarsal segments of
hind legs with conspicuous ventro-la’teral row of small spines.
Tibiae of fore legs with a long ventral spine at the apex. Tibiae
and first two tarsal segments of middle legs with small, paired,
apical spines; first tarsal segment bears also several similar spines
on its ventral border. Claws equal, about half the length of the fifth
tarsal segment, bifid. Empodium absent. Abdomen dark brown.
3 < 5 o
Hypopygium (fig. 17). Ninth segment-, sternite reduced to a
narrow strip of chitin; tergite not highly chitinised, bearing
apparently only two long bristles dorsally near its middle, the
chitinisation of the tergite interrupted a little posterior to them, the
posterior margin with a short hairy process on each side, the ventral
surface thickly covered by short hairs, the lobe-like processes well
developed, covered by short hairs, and bearing one or two longer
bristles. Forceps-, side-pieces rather long and narrow; claspers
short, terminating iri strong, pointed hooks, basal two-£hirds hairy.
Fig. 17. Spbacromias litoraurea , tp.n., outlines of male hypopygium. a , b, and c—
ventral views; d--dorsal view, a —forceps (bristles and hairs of tide-piece not shown) j
fr—aedoeagus; c —harpes 5 d —ninth tergite (small surface hairs not shown).
Harfes dark brown, highly chitinised, with a double dorso-ventral
curve; distal extremities fused to form a blunt, rather broad, process.
Aedoeagus conical, proximal portions of the limbs narrow and
highly chitinised, distal portions less highly chitinised; distal
extremity broad, with a slight ventral lip; ventral wall slightly
chitinised to about the level of the middles of the highly chitinised
portions of the limbs; membrane joining the aedoeagus to the ninth
sternite not spiculated.
Gold Coast : Odorkor, a small village near Accra, November,
1920; one male, obtained from a drain situated near a stand-pipe.
Weshiang, near Accra, June, 1921; one male, reared from plants
of the water-weed Pistia stratiotes, taken from the river Densu.
j6i
Genus BEZZIA , Kieff.
The chief characters of this genus, according to Kieffer (1919),
are—eyes glabrous, the last three or four antennal segments in the
male elongated, the wings bare or covered by microscopic setae, the
first and third veins not united by a cross-vein and forming a single
cell, the bifurcation of the fourth vein scarcely proximal to or under
the cross-vein, the femora on the fore legs at least armed with one
or more ventral spines in both sexes, the fourth tarsal segment
cordiform, the fifth unarmed in both sexes, and the claws small*
not half the length of the fifth tarsal segment, simple in both sexes,
or sometimes with a small median tooth in the female. Up to the
present we have not collected at Accra any specimens referable to
this genus, but we have received from Lagos a single female of one
species, and of this a description is given here.
Bezzia foyi , sp. n.
Measurements.
Length of body (one female) .2*1 mm.
Length of wing .1*4 mm.
Greatest breadth of wing..o’S mm.
Head dark brown, clothed with rather short dark brown hairs.
Eyes smooth, narrowly separated. Clypeus, proboscis and palpi
dark brown. Mouth-parts well developed, mandibles particularly
highly chitinised, and bearing strong teeth on their inner edges.
First palpal segment small, second short and broad, third, fourth,
and fifth sub-equal, about twice as long as broad; the third segment
scarcely at all inflated and without a sensory cup, but bearing
a patch of sensory hairs on its inner aspect; the fifth segment
pyriform, its distal end broad and rounded, bearing a few rather
long hairs. Antennae dark brown, the terminal segments rather
paler at their bases; hairs short, dark brown. First segment small:
torus sub-spherical, bearing numerous short hairs; third rather
longer than the fourth, with a short stem; segments four to ten oval,
slightly constricted at their bases, their lengths varying from once
and a half to twice the greatest breadth; segments eleven to fifteen
elongate, about three or four times as long as broad, the last segment
the longest and ending in a conical tip without a stylet. Thorax
3 6z
uniformly very dark brown, clothed with short, dark brown hairs,
and bearing above the wing bases a few longer, strong hairs.
Pleurae dark brown. Scutellum dark brown, not so dark as the
dorsum; bearing two lateral and two centro-marginal bristles and
very numerous short hairs. Post-scutellum dark brown. Wings
clear, slightly infuscated near the anterior borders, the stronger veins
brownish. Venation as shown in the figure (fig. 18 a). The costa
does not extend as far towards the tip of the wing as in the female
of Probezzia pistiae (p. 365); the fork of the fourth vein under
the cross-vein. The surface of the wing covered by microtrichia
and devoid of longer decumbent hairs; fringe very short on the
apical third of the wing. Halteres with pale brown stems and dark
brown knobs; knobs bearing a few short hairs. Legs dark brown,
conspicuously banded. Femora dark brown, especially those of the
a
Fig. 18. Bezzia foyi, ip.n. a —venation of wing of female (x 50 circa.);
b —ipermatheca (x 210 circa.)
•
hind legs, slightly paler basally, and in the case of the fore and
middle legs with a narrow pale band near the apex; knees dark
brown; tibiae dark brown, with a narrow pale band near the base,
and a less distinct, pale, sub-apical band; tarsi paler, last two
segments infuscated. Femora not swollen; fore femora armed with
two short, dark, ventral spines on its apical third, middle and
hind femora unarmed. Tibiae unarmed, not swollen. First tarsal
segment about twice as long as the second on the fore legs, relatively
longer on the middle and hind legs; bulbous hairs conspicuous on
the hind tarsus, forming two rows on the first and second segments,
and one on the third, on the middle legs are single rows of
similar hairs on the first and second segments. Fourth tarsal
segment cordiform; fifth unarmed. Claws simple, equal, short,
less than one-half the length of the fifth tarsal segment. Empodium
3*3
rudimentary. Abdomen dark brown, venter paler than the dorsum,
scantily clothed with short dark brown hairs. Spermathecae two,
highly chitinised, unequal, oval or egg-shaped and slightly
constricted sub-kpically (fig. 18 £); lengths about 103/1 and 84/1 in
the single female examined, and greatest breadths 53/* and 49/1
respectively; the duct narrow (about 4 p), and chitinised for some
distance (about 25j»).
NIGERIA (Southern provinces): Lagos, July, 1921 (Dr. H. Andrew
Foy); a single female taken in the evening upon the white lining of a
lamp-shade. We have pleasure in dedicating this species to the
collector, Dr. H. Andrew Foy, to whom we are also indebted* for
numerous other specimens of Ceratopogoninae from Lagos.
Genus PROBEZZIA , Kieff.
The chief generic characters of Probezzia, according to Kieffer
(1919), are—eyes glabrous, last four antennal segments in the male
elongated, wings bare or covered with microscopic setae, first and
third veins separated for their entire length, bifurcation of the fourth
vein scarcely proximal to or under the cross-vein, femora unarmed,
fourth tarsal segment cordiform, fifth unarmed in both sexes, and
claws small, about one-third the length of the fifth tarsal segment,
equal and simple in both sexes, or with a small median tooth in the
female. The two species which we have assigned to this genus
differ slightly in two respects from the above description, namely, in
having only the last three antennal segments definitely elongated,
the twelfth segment being but slightly longer than the eleventh, and
the fourth tarsal segment short and broad but not definitely
cordiform. These differences are very slight, or may even
depend on the manner of interpretation of the terms elongated and
cordiform.
Both the species described here were reared from plants of the
water lettuce Pistia stratiotes, but we were successful in procuring
for examination the early stages of the first species only. The larvae
are slender, almost white, eel-like organisms similar to those of
Culicotdes, but larger and with relatively longer and narrower heads.
They resemble the figure of the larva of Palfomyia longipennis
364
given by Malloch (1915). They appear normally to inhabit the
basal portions of the roots of the Pistia plants, and w6re reared to
the adult stage from plants taken from the water and brought, or
sent through the post, to the laboratory, and subsequently kept
merely moist. They are, however, capable of leading an aquatic
existence, and move very rapidly in water, swimming about in a
manner similar to the larvae of Culicoides. At the posterior end of
the body are long, stout hairs, which are an aid to progression. The
pupae are similar in form to those of Culicoides. They are able to
survive in water, but when placed in it quickly make for the side
and wriggle themselves above the surface. In the latter situation
they remain practically sedentary if undisturbed. The duration of
the larval stage was not determined, that of the pupal stage was
two to four days.
Probezzia pisiiae , sp. n.
Measurements. Male. Female.
Length of body .1*6 mm. 2'5 mm.
Length of wing .n mm, 1*9 mm.
Greatest breadth of wing.0*4 mm. o*6 mm.
The male is a much smaller, and much darker brown insect, than
the female. Head : occiput dark brown, with brown hairs. Eyes
glabrous, separated in both sexes. Clypeus and proboscis brown,
with brown hairs. Palpi dark brown, rather slender; the first
segment short, segments two, three and five sub-equal, about twice
as long as broad, the fourth segment rather shorter; third segment
not inflated, without a sensory cup but with a few sensory hairs
situated distally on its inner side. Mouth-farts somewhat similar
to those of midges of the Genus Prionognathus. Labium soft and
hairy. Labrum rather strongly chitinised, the proximal two-thirds
broad, the distal third tapering, fringed with delicate hair-like
processes. Hypo-pharynx broad, tapering disfally to a rounded
apex, and fringed. Mandibles (fig. 21 d), in the female, similar to
those of P. marmoratus , and similarly situated, but without teeth
and with only a few delicate hair-like processes on the outer edge;
the teeth on the inner edge are seven, large and strong, and proximal
to them is a row of delicate hair-like processes; in the male,
mandibles smaller, less highly chitinised and without strong teeth,
but with about five delicate hair-like processes on the inner side.
Maxillae rudimentary. Antennae : dark brown, the last five
segments in the female, and the last three in the male, darker than
the rest. First segment small, bearing a few short hairs in the
female. Torus yellowish-brown in the female, dark brown in
the male; bearing a few short hairs. Flagellum segments sub-
cylindrical : in the female, segments four to ten from twice to nearly
two and a half times as long as broad, segments eleven to fifteen
elongate, from nearly five to seven times as long as broad, the
fifteenth segment being the longest and ending bluntly; in the male,
the twelfth segment about two and a half times as long as broad,
the last three segments elongate, from three to five times as long as
broad, the fifteenth segment being the longest and ending bluntly.
Hairs short and scanty in the female; plume of the male poorly
developed. Thorax : dorsum uniformly dark or darkish brown,
clothed with short dark brown hairs and bearing later all y and
posteriorly a few dark brown bristles. Pleurae darkish brown.
Scutellum darkish brown, bearing a transverse row of six to eight
bristles, and in the female numerous scattered short hairs; in the
male are usually only six bristles, and the hairs are more scanty.
Post-scutellum dark brown, without a pit. Wings unspotted.
Venation as shown in the figures (figs. 19 and 20 a); costa in the
3^6
female extending further towards the tip of the wing than in the
male. Surface of the wing closely covered with microtrichia; no
macrotrichia present. Halteres pale brown, with dark brown knobs.
Legs almost uniformly infuscated in the male; in the female, more
or less banded, femora with a dark band before the apex, tibiae
with a dark apical band and a dark band a little beyond the base,
distal ends of first four tarsal segments and whole of fifth infuscated.
Femora not inflated and without strong spines. Tibiae moderately
hairy; fore and middle legs with a dark apical spine. Tarsus with
first segment more than twice as long as the second, fourth short
and broad, almost (especially in the male) but not definitely
cordiform, fifth unarmed. In the female, the first two tarsal
segments of the hind legs with a double row of ‘bulbous* hairs,
third segment and first, second, and third segments of the middle
legs with only a single row. Claws equal, short, not more than half
the length of the fifth tarsal segment; in the female, each with a
small but well developed basal tooth, in the male, smaller, bifid at
the tip. Empodium small, hair-like. Abdomen brown, clothed with
short dark brown hairs, which are most numerous cm the posterior
segments; in the female, eighth stemite bearing two highly chitinised
plates, one on each side of the vulva, clothed distally with numerous
short hairs. Spermathecae two, moderately chitinised, sub-spherical,
unequal, in one instance measuring 57 n by 50and 46/1 by 40^,
respectively; commencement of duct chitinised for only a short
distance (about 4/1 to 70).
Hypopygium (fig. 21 a-c). Hypopygium highly chitinised and
Fig. 2i. Probezzia pistiae , gp.n. a to c —male hypopygium. a —ninth tergite ;
b —harpes; c —forcepi and aedoeagus. i —mandible of female.
3 <n
dark brown, rotated so that the forceps lie dorsally .* Ninth segment :
tergite short, bearing at its posterior end two large, hairy, lateral
lobes, between which the spiculated lining membrane of the tergite
projects as a blunt process; stemite deep, slightly excavated in the
middle. Forceps : side-pieces short and broad, about as broad as
long, tapering slightly distally; claspers reduced to a small knob
bearing a few long hairs. Harfes highly chitinised, fused in the
middle line to form a stout chitinised rod, with a rounded, somewhat
expanded distal extremity. Aedoeagus highly chitinised, tapering
gradually to a narrow distal extremity; membrane joining the
ventral wall of the aedoeagus to the ninth stemite studded with
spicules.
PUPA. Length, female, about 3 mm., male, considerably smaller,
about 27 mm. Well chitinised; male very much darker than
female. Respiratory trumpets usually bent posteriorly, rather short
and broad, length about 0*2 mm:, ratio of length to breadth about
7 to 1, smooth and without knob-like processes, the distal end
infuscated. The main tracheal trunk is broad, devoid of lateral
branches, and terminates in a number (about eighteen) of short,
blunt processes which lead to the surface and are arranged in the
form of an inverted U. Cephalo-thorax of the usual form, but the
separation of cephalic and thoracic portions is ill-defined. Anterior
marginal tubercle very small, bearing a long bristle; above the
highest part of the antennal case (the beginning of the flagellum
part) are two socket-like marks, apparently unarmed; a little further
back, and in front of the base of the trumpet, is a small tubercle
bearing a minute spine; anterior dorso-median tubercle very small,
bearing two small sockets, the inner unarmed, the outer bearing a
short, spine-like hair, dorso-lateral situated very close to the base
of the trumpet, rounded and irregular in form, bearing a long and
a shorter hair and an apparently unarmed socket; ventro-lateral
ill-defined, bearing a short hair and an unarmed socket; ventro-
median represented by two moderately long hairs. Dorsal tubercles
much reduced : anterior double, each half bearing a moderately long
hair, lateral bearing a similar hair, posterior bearing a rather
smaller one. Anterior to the dorsal tubercles is a transverse row of
# It will be convenient, however, for descriptive purposes to continue to refer to the surface
on which Hes the tergite as the dorsal, and to that on which lie the stemite and the aedoeagus as
the ventral.
368
highly chitinised rugae, and posterior and external to the lateral
tubercle is an unarmed socket. Postero-dorsal tubercle obsolete.
Posterior margin of dorsum rounded, not extended backwards as a
median process. Abdomen : first segment small and narrow, second
large and broad, the others decreasing progressively in breadth
towards the apex. Integument spiculated, especially in the male,
and with pigmented areas similar to those of Stilobezzia spirogyrae .
Anal segment with acutely pointed, slightly divergent, dark-tipped
processes. Dorsal tubercles: antero-submarginal, small, situated
close together and almost contiguous, each bearing a hair, the
outermost the larger; postero-marginal, four, the inner small, bearing
a minute hair, the next merely an apparently unarmed socket, the
outer two larger, contiguous, each bearing a hair. Ventro-lateral
tubercles: all arising from a central projection of the segment;
antero-submarginal, small, bearing a hair, and a little dorsal and
slightly posterior to this tubercle an apparently unarmed socket;
postero-marginal, two, well developed, the ventral bearing a hair,
the dorsal double and bearing two hairs. Ventral tubercles:
postero-marginal, three, almost contiguous, the inner bearing a short
hair, the other two longer hairs.
Larva. The larva is eel-like and slender, pale coloured or
nearly white; length 7 mm. to 9 mm., greatest breadth about o*2 mm.
to 0 3 mm. Head yellowish-brown, long and narrow, length about
04 mm., greatest breadth about 01 mm Eyes black, bilobed or
reniform, situated laterally a little anterior to the middle of the
head. Antennae and palpi small. Hairs quite small, and some
apparently unarmed tubercles also present: on the dorsal surface,
one pair anterior admedian situated about the level of the
mandibles, two pairs (the more anterior with a divided hair)
and a pair of small, apparently unarmed, tubercles anterior dorso¬
lateral situated at about the level of ihe comb-like part of the
hypopharyngeal sclerite, one pair of apparently unarmed tubercles
central dorso-lateral, two pairs posterior dorso-lateral, and two pairs
bearing small spines posterior admedian at the extreme posterior end
of the head; on each lateral surface, one anterior, two about the level
of the eye, and a small tubercle, apparently unarmed, a little more
posteriorly; on the ventral surface, two pairs anterior admedian, and
one pair central ventro-lateral. Mental plate with a strong, pointed,
3<*9
central tooth, and two more delicate teeth on each side. Hypo-
pharynx not very strongly chitinised, the posterior sclerite comb-like,
bearing about a dozen pointed teeth. Mandibles large and highly
chitinised, base expanded, distal portion a powerful hook. Body
cylindrical, composed of twelve elongated segments each bearing a
few minute hairs. On the distal end of the anal segment are fourteen
stronger hairs arranged as follows: dorsally and ventrally two pairs
of long, stout, dark hairs about half the length of the anal segment,
and two shorter hairs; laterally, on each side a single short hair;
these hairs are usually turned anteriorly, and ip life appear to be of
assistance in progression. Anal gills of the usual form, rather short,
being about one quarter the length of the anal segment, deeply cleft
into two pointed processes.
Gold Coast: Oblogo, near Accra, December, 1920, to June,
1921, numerous specimens reared from plants of the water lettuce
(Pistia stratiotes ), taken from a swamp and from the river Densu
(PI. XXI, fig. 1). The larvae, while quite capable of leading an
aquatic existence, appeared normally to frequent the roots of this
plant, and were frequently reared to the adult stage in plants kept
merely moist.
Probezzia stephensi , sp. n.
This insect, of which at present we possess only a single
male, resembles Probezzia pistiae , and, indeed, was originally
included among some examples of that species in our collections
on account of its almost identical colour markings. Subsequently
certain morphological differences were observed, particularly in the
hypopygium, which warrant its separation as a distinct species.
Only the chief differences between this species and P . pistiae are
here given.
Measurements.
Length of body (one male) .17 mm.
Length of wing .To mm.
Greatest breadth of wing.0*4 mm.
Head (fig. 20 b) facets of eyes very widely separated (by about
105/1) dorsally; anterior median angle of the occiput broad and
obtuse. Thorax with rather fewer hairs on the dorsum, and some
37 °
of them larger than those in P. pistiae ; scutellum bearing two sub-
median and two lateral bristles, and a few (about a dozen) short
hairs. Legs : hind legs with a single row of ‘ bulbous ’ hairs on the
first tarsal segments only. Claws with terminal fork deeper.
Abdomen bearing shorter and more scanty hairs.
Hypopygium (fig. 22). Ninth segment : stemite reduced to a narrow
strip of chitin; tergite short, tapering, bearing only two large bristles
which are situated one on each side near the posterior margin,
continued posteriorly as two large processes, bearing hairs and a few
bristles, which are separated from each other in the middle, and with
a large, hairy, lobe-like process on its under surface. Forceps-.
Fig. 22. Probtxzia supbensi , ip.n., outlines of male hypopygium. a —ventral
view; b —lateral view, (x 250 circa.)
side-pieces of usual form, moderately hairy; claspers of usual form,
rather short (about half the length of the side-pieces), broad and
without large hairs at the base, narrowing abruptly in the middle,
and terminating in a somewhat spoon-like end. Harpes in lateral
view bent in the middle at a right angle (see fig. 22 b ) 9 the distal
halves somewhat beak-shaped; in dorso-ventral view the distal
halves appear as two contiguous chitinous plates (see fig. 22 a).
Aedoeagus broad, tapering, bearing on each side at the distal end
a barb-like process; ventral wall chitinised almost to the base;
membrane joining the aedoeagus to the ninth stemite devoid of
spicules.
4
37 1
Gold COAST: Oblogo, near Accra, March, 1921; reared from
some plants of the water lettuce (Pistia stratiotes ), taken from the
river Densu. (PI. XXI, fig. 1). We have much pleasure in
dedicating this species to Dr. J. R. C. Stephens, to whom we are
indebted for numerous interesting collections of biting midges from
the Ilorin Province, Nigeria.
Genus DICROBEZZIA , Kieff.
The chief characters of this genus, which otherwise resembles
Bezzia , according to Kieffer (1919), are—wings with the third vein
reaching as near to the wing apex as the anterior branch of the
fourth vein, femora unarmed, fourth tarsal segment cordiform, fifth
tarsal segment in the female armed with several pairs of black
‘batonnets/ in the male unarmed, and the claws in the female
equal, bifid, the large branch two-thirds the 'length of the fifth
tarsal segment, the small branch two-thirds the length of the large
branch, in the male small and simple. We have referred the
following species to this genus largely on account of the fact that
the fifth tarsal segment in the female is armed with short, dark
spines, but it should be pointed out that in other respects it does
not entirely agree with the generic description given above. The
differences are considerable, and would, perhaps, warrant the
erection of a new genus.
The species closely resembles Probezzia pistiae , both as regards
the general morphology of the adults, including the structure of the
hypopygium of the male, and the characters of the pupa. In the
following description, therefore, only the chief points of difference
are noted.
Dicrobezzia nigritibialis, sp. n.
This insect, of which at present we possess only a single male
and a single female, resembles closely Probezzia pistiae ; the chief
differences between it and the latter species are as follows: —
Measurements. Male. Female
Length of body .2*3 mm. 27 mm.
Length of wing . 17 mm. 2*2 mm.
Greatest breadth of wing.0*4 mm. o‘6 mm.
The female is darker than the male. Head very dark brown.
Eyes separated in both sexes. Palpi in the male small, stumpy,
tapering distally, all the segments very short; in the female, longer,
as in P. pistiae, Antennae \ torus almost black in both sexes:
middle segments of the flagellum paler brown in the male, in the
female, antenna entirely dark brown. Thorax : dorsum uniformly
dark brown, almost black; hairs scanty. Pleurae very dark brown.
Scutellum very dark brown, bearing in the male, two sub-median and
two lateral bristles and a few short hairs; in the female four sub¬
median and two lateral bristles and numerous short hairs. Wings very
delicate and with a white appearance, venation as in P. pistiae ; micro-
trichia extremely small and delicate. Halteres with pale yellowish-
brown knobs, and darker stems. Legs : femofa and tibiae very dark
brown, first four tarsal segments pale, slightly infuscated at their
apices, last tarsal segment entirely dark. Fourth tarsal segment
in the male longer * than in P . pistiae , cylindrical; in the female,
shorter, especially on the fore legs. Claws of the male small, equal,
bifid at the tips; those of the female large, each with a well developed
basal tooth. Fifth tarsal segment in the female armed with
numerous (twelve on the fore legs, fourteen on the middle and hind
legs) strong, black spines; in the male unarmed. Abdomen dark
brown, but not so dark as the thorax, and in the female paler
proximally; in the female, eighth stemite bearing a few long,
stout hairs on each side of the vulva. Spermathecae two, highly
chitinised, sub-spherical and unequal (diameters about 76 fi and 60/1 L
respectively); commencement of the duct chitinised for a short
distance (about 10 fi).
Hypopygium (fig. 23). Hypopygium very dark and highly
chitinised. Ninth segment : stemite deep, excavated in the middle
posteriorly; tergite not very highly chitinised, bearing at its
posterior end two prominent, hairy, lobe-like processes, spiculated
portion of its lining membrane prominent. Forceps : side-pieces
moderately developed, hairs not very long, distal extremity conical;
claspers obsolete. Harpes very densely chitinised, fused in the
middle line, and projecting backwards as a process which does not
expand at its end, but appears to be double; in dorso-ventral view
the posterior projection appears to be straight but not all in the
same focus, in lateral view it is seen to be bent sharply towards the
373
tergite near its distal end. Aedoeagus : form somewhat similar
to that of P. stephensi , but more densely chitinised. Membrane
joining the aedoeagus to the ninth stemite studded with spicules.
PUPA. The pupa is very highly chitinised, especially that of the
female, and coarsely spiculated. Length, male 3'6 mm., female
4'3 mm. It differs chiefly from the pupa of P. pistiae in the
following points. Respiratory trumpets not turned backwards,
relatively shorter and broader than those of P. pistiae, terminal
branches of the tracheal trunk only about ten in number. Cepkalo-
thorax : dorso-lateral tubercle bearing, apparently, only a single
Fig. 23. Dicrobezzia nigritibialis , sp.n., outlines of male hypopygium. a —ventral
view of forceps and aedoeagus; b —lateral view, middle focus showing harpes and
aedoeagus. (x 190 circa).
hair, ventro-lateral composed of two small nipple-like processes,
each armed with a stout hair; ventro-medisln apparently represented
by a very minute hair. Dorsal tubercles very small: anterior
double, the one part anterior to the other, each bearing a minute
spine; lateral bearing a long hair; posterior bearing a minute spine.
Abdomen : cuticle coarsely shagreened; large, dark, lateral macules
at the anterior margin of the segments dorsally, in addition to the
macules noted in P. fistiae . Anal segment: cases for forceps
small and dark coloured. Dorsal tubercles : antero-submarginal not
contiguous, a small hump with a socket-like mark posterior to the
outer one; postero-marginal, three, the innermost small, bearing a
small spine, the other two situated much more laterally, the inner
rather large, bearing a short spine and having on its inner side an
374
unarmed socket, the outer small, bearing a hair. Ventro-lateral
tubercles: antero-submarginal two, the dorsal one bearing a small
spine, the ventral a hair; postero-marginal, two, well developed,
each bearing a small spine. Ventral tubercles, three, practically
contiguous, the inner bearing a short spine, the middle a long hair,
and the outer a short hair.
Gold Coast: Weshiang, near Accra, June 29th, 1921; two
pupae found in a sample of the algae growing in one of the
reservoirs of the Accra waterworks. We are indebted to Mr. R.
Simmons for bringing these specimens to the laboratory.
REFERENCES
Carter, H. F., Ingram, A., and Macfix, J. W. S. (1920 and 1921). Observations on the
Ceratopogonine Midges of the Gold Coast. Puts I to IV. Annals Trap. Med.
Parasit ., Vol. IV, pp. 187-210, 211-274, 309-331; Vol. V, pp. 177-212.
Knrm, J. J. (1913). Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientak (191L-1911).
Insecies Diptbes J. Cbironomidae et Ceeidomyidae. Paris.
-(1917). Chironomides d’Amtrique conserves au Mus6e National Hongrois de Budapest
Ann. Mus. Nat. Hung., Vol. XV, pp. 292-364.
- (1918). Chironomides d*Afrique et d*Asie conserves au Musle National Hongrois de
Budapest. Ann. Mus. Nat. Hung., VoL XVI, pp. 31-136.
- (1919). Observations sur les Chironomides (Dipt.) d6crits par J. R. Malloch. Bell.
Soc. Ent. France, No. 10, pp. 191-194.
- (1919). Chironomides d’Europe conserves au Mus£e National Hongrois de Budapest
Ann. Mus. Nat. Hung., Vol. XVII, pp. 1-160.
Malloch, J. R. (1915). The Cbironomidae or Midges of Illinois with particular reference to
the species occurring in the Illinois River. Bull. III. State Lab. Nat. Hist., Voi. X,
Article VI, May.
376
EXPLANATION OF PLATE XXI
Fig. i. The river Densu at Oblogo, near Accra, showing the
Pistia plants from which were reared the following
midges:— Culicoides distinctipennis, Dasyhelea incon-
spicuosa, Prionognathus pseudotnaculipennis , Ketnpia
ochrosoma, Probezzia pistiae , and P. stephensi.
Fig. 2. Pool at Accra, from the marginal mud of which were
reared the following midges: —Culicoides austeni %
C. distinctipennisy C. neaveiy C. schultzeiy C. similis ,
Dasyhelea fuscipleurisy D . inconspicuosa, and Stilo -
bezzia spirogyrae.
377
THE EFFECT OF SALINE SOLUTIONS
AND SEA-WATER ON STEGOMYIA
FASCIATA
BY
J. W. S. MACFIE
(Received for publication 24 August , 1921)
Stegomyia fasciata is relatively intolerant of salt (NaCl). In
some experiments carried out in 1915 it was found that the larvae
died rapidly in 2 per cent, salt solution, that the gravid adult
females were relucant to lay their eggs on this medium, and that if,
faute de tnieux , they did so the eggs were killed and did not even
harden and darken. Moreover, it was found that normal eggs of
5 . fasciata placed in salt solution of about the same strength
(2*3 per cent.) failed to produce living larvae. From these and some
previous observations (1914) it was thought that sea-water might be
found to be of service in the campaign against this mosquito.
More recently these experiments have been repeated at Accra
actually using sea-water. It was found that undiluted sea-water
killed the larvae in a few hours (two to four), and that when diluted
with tap-water 50 per cent, (equal to about 1*6 per cent. NaCl) or
over was fatal within twenty-four hours. As regards the influence
on the gravid females and the action on the eggs, the results were
similar to those obtained with salt solution as is shown by the two
following experiments.
Experiment I. 20th May, 1919. Stegomyia fasciata , two females and one male
in a jar containing sea-water. Females fed this day.
22nd May. Male dead.
23rd „ No eggs. Females fed at 10 a.m.
25th „ 4 p.m.—many eggs on the water, all white.
5th June. No larvae have hatched.
Experiment II. 20th May, 1919. Stegomyia fasciata y one male and one
female in a large jar containing sea-water in which stands a small beaker containing
tap-water.
23rd May. Female fed.
24th „ Male dead.
27th „ Many eggs on the tap-water, a few only on the sea-water : the
latter are white.
1st June. Many larvae in the beaker containing tap-water, none in the
sea-water.
378
Sea-water, in fact, was found to act in a similar manner to a
solution of common salt of equivalent strength.
An attempt was made to determine the highest percentage of salt
that the larvae could tolerate. Such experiments are not .easily
devised because normally the larvae pupate after a few days, and it
is therefore only possible to determine the percentages which prove
fatal rapidly. Two series of experiments were carried out with the
object of obtaining information on this point. In the one series
larvae were used which were in the state of arrested development,
to which reference has been made elsewhere (1915), and showed no
inclination to pupate. Such larvae were placed in jars containing
100 c.c. or 50 c.c. of water to which each day o*i gm. of salt was
added in a 10 per cent, solution; at the end of each experiment the
amount of salt present in the medium was determined by titration.
In four such experiments in which the strength of the saline medium
was increased by o* 1 per cent, daily, the larvae did not survive more
than 0*8, 09, 077, and 0*87 per cent. NaCl respectively. In four
other experiments in which the strength of the saline medium was
increased by 0 2 per cent, daily, they did not survive beyond
r6, r8, 09, and ri per cent. NaCl respectively. The results of
these experiments were not quite satisfactory because in the control
jars containing only tap-water some of the larvae died, showing that
either the larvae in this state were delicate or that they were suffering
from the lack of suitable or sufficient food.
It was thought, therefore, that a better practical test of the
amount of salt tolerable to the larvae of S. fasciata .would be obtained
by starting a culture of these insects in a medium containing a
low percentage of salt, allowing it to concentrate naturally by
evaporation, and noting the point at which the larvae died. Such
experiments it was thought would also show if the larvae were able
to become habituated to high degrees of salinity.
An experiment was, therefore, started with a natural medium,
rich in organic material, in a large jar on the sides of which were
very many ripe eggs of 5 . fasciata . Sufficient common salt had
been added to the medium to bring the percentage of NaCl up to ri.
The jar, covered only by a piece of gauze, was then placed on the
laboratory bench and allowed to concentrate gradually by natural
evaporation. The larvae which hatched from the eggs developed
379
rapidly at first, then more slowly, and after about a fortnight
appeared to be dying off. On the nineteenth day, when it was clear
that they were rapidly diminishing in number, a small quantity of
the medium was withdrawn for analysis. It was found to contain
i "3 per cent. NaCl. From this time onwards the larvae steadily
dwindled, the last individual dying on the thirty-sixth day of the
experiment, when the salinity of .the medium was found to be
l *45 per cent. NaCl. During the experiment only a very few of
the larvae pupated, and all that did so, excepting the first (which
pupated on the ninth day, when the salinity was estimated to be
i'2 per cent.), died in the act. A second experiment on the same
lines may be summarised as follows: —
i»t Day. Salinity of medium equals i*i per cent. NaCl. Multitudes of little
larvae which have just hatched.
8th „ Larvae fewer and not growing much.
i6th „ Very few surviving larvae. No pupae yet.
18th „ Only two surviving larvae.
20th „ Last larva dead. Salinity of the medium found to be 1*38 per cent.
NaCl. No pupae have appeared.
The four experiments of this sort (see table) that were carried
out showed that the percentages of salt in the media at the times
of the deaths of the last larvae were 1*45, 1 *38, 145, and 1*45,
Number of days
for which the
experiment lasted
Percentage of NaCl
in the medium at
the beginning of the
experiment
Percentage of NaCl
in the medium at
the end of the
experiment, namely,
when the last larva
died
Number of pupae
36
i*i
r 45
1, and a few which died.
20
1*1
1*38
None.
*4
ro
r 45
None.
r 45
respectively. The larvae had, of course, begun to die off some
considerable time before this concentration was reached. Pupation
was very seldom attempted, and was usually fatal. The experi¬
ments furnished no evidence that the larvae could be habituated to
such degrees of salinity.
380
.The inference from these experiments would appear to be that
a ro to 1*4 per cent, solution of common salt, or an equivalent
strength of sea-water, would effectually prevent the larvae of this
mosquito from developing to the adult stage. It would seem
probable that sea-water, if used for such purposes as flushing drains
and gutters, scowering market-places, etc., would kill both the
larvae and the eggs of 5 . fasciata , and that even if puddles were
left the adult females would be reluctant to deposit their eggs on
them, but that if they did so the eggs would be killed immediately.
REFERENCES
Mactix, J. W. S. (1914 and 1915). Bull EnU Res., IV, p. 339, and VI, p. *25.
*
3 *i
THE PREVALENCE AND CHARACTER
OF TUBERCULOSIS IN HONGKONG
BY
HENRY HAROLD SCOTT,
M.D., M.R.C.P. Lond., F.R.S.E., D.P.H.
GOVERN MINT BACTKKIOLO GIST, HONGKONG;
LECTURER ON SPECIAL PATHOLOGY, HONGKONG UNIVERSITY
(Received for publication 12 October , 1921)
III. THE MORBID ANATOMY AS MET WITH IN CASES
AMONG CHILDREN
It is a well-known fact that the primary portal of entry of the
bacilli in cases of tuberculosis and the mode of spread of the disease
are by no means always easy to determine: in some instances,
indeed, one can hardly do more than hazard a conjecture. When
we remember that the bacilli may pass through a mucous membrane
and even through the walls of vessels and circulate as foreign
bodies without setting up any immediate injury, but only more
remotely causing changes at some distant site where they finally
settle, we must always be cautious against employing too freely the
anatomical distribution of the lesions as found in the post-mortem
room for the interpretation of their genetic relations. In some
instances, again, none of the ordinarily described routes seem to
explain the method of spread, as was indicated in some of those
mentioned in a previous paper.
The extent and distribution of tuberculous lesions in an animal
inoculated experimentally depend upon several factors, namely, the
number and virulence of the bacilli, the resistance set up against
infection by the inoculated animal, the seat of inoculation, and the
time which has elapsed since infection. It was found as a result
of several experiments in which the same dose of bacilli from the
same source was inoculated at the same site (namely, subcutaneously
into the left hind leg) into animals of the same species and as nearly
as possible of the same weight, that in ten days the adjacent gland
was involved; in ten to twenty days the left superficial and deep
382
inguinal and the sacro-lumbar glands, and also, perhaps, the spleen
and the retrohepatic glands. In another ten to fifteen days the
liver, the lungs, the bronchial, suprascapular, and cervical glands
on both sides showed involvement (Delepine).
Since in my series the respiratory portal of entry was that most
frequently encountered, these cases will first be dealt with.
Albrecht, Ghon, and others hold that there is a special form of
tuberculosis in children, consisting of a primary lung focus and
resulting from the entrance of the bacilli by inhalation. This focus,
they stated, was usually the size of a pea, but might be quite small
and was rarely larger than a cherry. In nearly three-fourths of the
cases the focus was single. The focus is believed to arise
* in aggregations of lymphoid tissue in the neighbourhood of small
bronchi/ Around this focus small tubercles are seen, and perhaps
reactionary fibrous tissue. In the case of larger foci, hard, dry
caseation is usual, and occasionally there is actual cavitation. As
regards the situation of the focus, the order of frequency was: right
upper, left upper, right lower, left lower, right middle, the first-
named site being four times as common as the last.
An examination of the mediastinal glands showing involvement
will, in many cases, enable one to predict the situation of the lung
focus. The lymphatics, deep and superficial, discharge into the
broncho-pulmonary glands situated between the branches of the
main bronchi and at the hilus; those from the middle and lower
portions of the lung into the inferior tracheo-bronchial at the
bifurcation of the trachea; those from the upper into the superior
tracheo-bronchial in the angle between the trachea and bronchus.
There is also a chain of glands each side of the trachea, the para-
tracheal glands. Infection across from one side to the other is
frequently met with.
The original focus may open into a bronchus, and thus by
inhalation a tuberculous broncho-pneumonia is set up ; a like result
would, of course, follow the perforation of a bronchus by an inherent
gland. Again, there may be direct extension from a gland adherent
to the pulmonary tissue, while, lastly, by communicating with a
blood-vessel, miliary tuberculosis may ensue.
It used to be held that gland infection was primary and the lung
condition secondary to it, but this would leave unexplained the
3«3
fact of apparently arbitrary selection of a site remote from the
primary gland infection, while the intermediate tissue remained free
from disease.
Canti examined the bodies of eighty-four children under ten
years of age. Of these, there were thirty-three under one year, and
sixteen (19*05 per cent.) showed tuberculous lesions. Of these
sixteen, ten had foci in the lungs, the largest was the size of a
cherry, the average that of a pea. In eight a single focus only was
found; in one case two foci of practically the same age were seen,
and one showed several; in this instance, however, one of the foci
was cavernous and appeared older than the remainder. In the
eight with a single focus, this was found three times in the left lower
lobe, twice in the right upper, twice in the left upper, and once in
the right middle. In other words, the findings in this series of
Canti’s agreed in the main with those of Ghon as affording evidence
in favour of the common existence of pulmonary tuberculosis in
children. The chief points to which attention is directed are the
following, and, in discussing the series of cases considered here,
I do not think one can improve upon the lines taken by Canti, and
for purposes of comparison it is advisable to deal with the points in
this order: —
1. The almost constant finding of a lung focus when tuberculous
mediastinal glands are present, and the close relation of
these glands to the lung focus.
2. The frequent singleness of the lung focus.
3. The constant finding of tuberculous mediastinal glands when
a lung focus is present—a corollary of the first.
4. The almost constant absence of a lung focus when the portal
of entry appears to be elsewhere.
5. The almost constant absence of evidence that the portal of
entry .may be elsewhere when a lung focus is present—a
corollary of the last. h
Work at the mortuary here has afforded me exceptional
opportunities for studying these questions, the number of bodies to
be examined is great, the proportion of children very highland a
few weeks’ experience sufficed to drive home the fact that tuberculosis
forms a large percentage of the causes of death.
The differences between the post-mortem findings in children
3«4
dying from tuberculosis and those in adults are considerable.
Amongst the first three hundred consecutive cases with which this
and the two previous papers are concerned there were two hundred
and twenty-five under ten years of age. The remaining seventy-five
are insufficient for a study of adult tuberculosis, so this paper will
be restricted to dealing with the disease as it occurs in children
here. The number (two hundred and twenty-five cases) will
provide sufficient basis for argument as to whether the conditions
of tuberculosis in the tropics, as exemplified at least in Hongkong^
resemble those at home, and, if not, in what the differences consist.
For purposes of discussion, it will be well to take the points in the
above order.
I. The almost constant finding of a lung focus when tuberculous
mediastinal glands are present , and the close relation of these
glands to the lung focus.
The truth of this statement has been substantiated in the majority
of the present series. In twenty-nine instances, however, caseous
mediastinal glands were found without any focus being detected
in the lung9. Eleven of these showed strong evidence of being
primarily alimentary and the mediastinal glands may have become
involved secondarily to the mesenteric, a condition the occurrence
of which was proved by the experimental work of Calmette, Guerin
and Breton (1907). They found that in guinea-pigs dying in two
to four weeks after being fed on the bacilli the mesenteric glands
(especially the superior deriving from the small intestine) were
enlarged and inflamed, although no trace of any intestinal lesion
could be determined. After six to seven weeks, these glands were
caseous in greater or less degree and the lungs showed involve¬
ment by miliary tubercles with affection of the corresponding
tracheo-bronchial glands. Furthermore, these glands, as shown in
several of the cases detailed in this series, become caseous more
rapidly than the pulmonary lesions preceding their involvement.
In one other of my series there was a tuberculous abscess of the
sixth cervical vertebra discharging into the right pleural cavity,
which might account for the involvement of the mediastinal glands
without a focus in the lung. Putting these aside, there were still
seventeen which did not conform to the statement relative to the
385
presence of a lung focus when mediastinal glands are found and the
relation of the glands to the focus. Of these seventeen, there were
fifteen which showed miliary tubercles in the lungs, occasionally in
considerable numbers, but in twelve only a few; nevertheless,
mediastinal glands on one or both sides were found enlarged and
caseated. In none of them was any sign of tuberculosis found in
the tonsils or cervical glands. Where the involvement of the lung
with miliary tubercles is fairly extensive* it may be argued that the
gland constituted the focus whence the lung became infected, but
then we are still in the dark as to the source whence it became itself
tuberculous.
Apropos of some of these cases, the remarks of Bushnell in the
Military Surgeon (1918) may be recalled. He states that from a
hilus infection the tubercle bacillus is described as travelling by the
peribronchial lymph spaces in a direction opposite to that of the
normal lymph flow to a region in the upper lobes where the lymph
motion is most sluggish. This might be aided by a reversal of the
lymph current, and such a reversal might in turn occur as the result
of a block at the hilus. The tubercle bacilli travelling by these
spaces to the parenchyma are resisted by the tissue cells and a
type of peribronchial tuberculosis results. Caseation through the
bronchus may take place, although peribronchial tuberculosis is
more often of the 1 closed ’ type, as is evidenced clinically by the
frequency with which the bacilli are found in the sputum and the
rarity of haemorrhages in such cases.
In two others the difficulty is increased by the fact that although
the mediastinal glands were enlarged and caseous, in one case the
tracheo-bronchial, in the other the paratracheal, forming adherent
and caseous masses, there was no involvement of the lungs at all.
In the former there were meningeal tubercles mostly at the base and
along the sylvian fissures, whereas in the latter the only organs
found affected were the kidneys, where at the base of a pyramid
towards the lower pole of each were several minute tubercles focally
arranged.
So much for instances in which caseated mediastinal glands
were found without a corresponding focus in the lung. As regards
the second point—the close relation of the affected glands to
the lung focus—there were four cases in which the two did not
386
correspond, in other words, the expectation of localising the lung
focus from the gland involved was falsified. In three the focus
was in the left lung, as large as a haricot bean, and in one case
there was a cavity as large as a filbert; in the fourth the focus was
in the right lung. In each case there was a caseated gland, but on
the opposite side. The related glands on the affected side were,
in three, not involved at all, and in the fourth there was a little
congestion only.
It is well known that communication between the glands from
one side to the other may be free, and this may be offered as an
explanation of the passage of infection from one side to the other;
nevertheless, it is less likely that in children, in whom caseation of
the mediastinal glands occurs early and readily, infection should
pass to the opposite side and apparently miss those on the side
affected. One example may be quoted briefly:—In a boy, three
years of age, there was a sub-apical focus as large as a pea in the
right lung and a few scattered miliary tubercles in both; the hilus
glands were enlarged on both sides, but whereas on the right they
showed merely a small caseous point on section, on the left the gland
was completely caseated. This differs from the four previously
mentioned in that there was a spot of caseation in the gland on
the side of the focus, whereas in the others the related glands had
apparently escaped altogether.
2. The frequent singleness of the lung focus.
Among the sixteen cases described by Canti there were eight
with a single focus, while Ghon states that in 72 35 per cent, this is
the case. Of the two hundred and twenty-five children under
ten years of age among my series of three hundred cases, there were
one hundred and thirty-seven showing focal conditions in the lungs.
Of these, there were ninety-five in which the focus was single,
i.e. t in 6934 per cent.; in forty-two, or 30*66 per cent., there was
more than one.
Ghon found that when only one lung focus was discovered the
various lobes were involved in the following order of frequency: —
Right upper 3098 per cent., left upper 23*24 per cent., right lower
22 54 P er cent., left lower 15*49 per cent., and right middle
7*75 per cent.
387
Of the ninety-five cases in my series in which only a single focus
was found, the numbers in which each lobe was concerned were: —
Right upper twenty-six, or 27*37 per cent.; right lower twenty-three,
or 24*21 per cent.; left upper eighteen, or 18*95 per cent.; left lower
fifteen, or 15 79 per cent.; right middle thirteen, or 1368 per cent.
The main differences, it will be seen, are that in his series the left
upper and right lower were affected in about an equal number of
times, the former slightly preponderating, whereas in mine these
were reversed, and, secondly, in mine the proportion in which the
right middle lobe was involved was much higher.
As regards those which contained more than one focus, there
were thirty-two with two, five with three, and the same number
with several. The following was the distribution in cases with
two foci: —
Both in the upper lobe of the left lung. 5
Both in the lower lobe of the right lung . 5
Both in the lower lobe of the left lung. 4
Both in the upper lobe of the right lung . 3
One each in the lowest and middle lobes of the right lung ... 4
One each in the upper and lower lobes of the left lung ... 4
One each in the upper and middle lobes of the right lung ... 2
One each in the upper and lowest lobes of the right lung ... 2
One each in the upper lobe of each lung . 2
One each in the lower lobe of each lung . 1
Of the five instances in which three foci were found, in one case
all were in the upper lobe of the right lung; in another all were in
the lower lobe of the left. Of the remaining three, all were in the
right lung, viz., one in the upper and two in the lower in two cases,
and the reverse of this, two in the upper and one in the lower
in one.
Finally, in the five cases in which there were more than three
foci, the following was the distribution:—All in the left lower lobe
in one; in another, three foci were present in the upper lobe of the
left and one in the upper of the right; in a third, foci were present
in all lobes except the right upper; in a fourth, in all except the
left upper; in the fifth, in all except the left lower.
1 Taking all the cases in which foci were found in the lungs,
t.e., in one hundred and thirty-seven instances, the right upper lobe
388
was involved forty-two times, the right lower forty-one, the left
upper thirty-two, the left lower twenty-eight, and the right middle
twenty-two. Thus, whereas Ghon found that the left upper was
that most frequently involved after the right upper, in my series
these were reversed; the right upper and right lower were involved
almost an equal number of times, whether we consider merely cases
with a single focus or whether we have regard to all instances in
which focal lesions were present.
3. The constant finding of tuberculous mediastinal glands where a
lung focus was present .
In adult cases this has certainly not been my experience here,
but it appears to hold good with nearly all cases in children. In
twelve of this series, in spite of focal affection of the lung, and even
of considerable advancement of the disease, there was no naked-eye
involvement of mediastinal glands. In four of these the glands
showed microscopically giant cells and a few bacilli. In three
others, owing to the early age at which death occurred (twenty-two,
twenty-four and twenty-nine days, respectively), although foci
were present and the disease was of considerable extent in the
lungs, an explanation of the absence of tuberculous affection of the
corresponding mediastinal glands may be tendered by suggesting
that death took place before there was time for gland involvement
to arise. This hypothesis finds a certain measure of support from
two others, each of seven weeks old, in whom there was a lung focus
and the corresponding mediastinal gland was congested and slightly
swollen, not macroscopically tuberculous, but on sections being made
giant cell systems and bacilli were seen. On the other hand, the
findings in two others appear to deprive this hypothesis of its value.
In one of these, a child of three years, there was typical phthisis-
tuberculous broncho-pneumonia with ulceration—and in the other,
four years of age, the disease had been in existence long enough to
produce caries of three vertebrae in addition to a lung focus
and several caseating tubercles; nevertheless, no corresponding gland
involvement was found.
3»9
4. The almost constant absence of a lung focus when the portal of
entry appeared to be elsewhere .
This is a more difficult matter on which to pass an opinion out
here, where the chances of infection by a double route, respiratory
and alimentary, are so great. In a previous paper, when we were
discussing cases in which the primary portal was uncertain, several
of such were dealt with. To take these as instances for considera¬
tion as to the verity of the dictum of this section would be to beg
the question. The statement would apply rather to places where
one sees either respiratory or alimentary cases, or at least cases in
which the primary portal is undoubtedly one or the other, not those
in which the dual route is not only possible, but, as in many here,
highly probable.
Apart from these, there were five in which a definite lung focus
was present, although the primary portal of entry was probably,
in fact, one might say certainly, not via the respiratory tract. In
two there was a condition cf tabes mesenterica, the glands in the
abdomen being in aggregated caseous masses; in one of these cases,
an infant of eight months, there was a focus the size of a pea in the
right lung; in the other, a child of twelve months, a focus as large
as a marble, also in the right lung. In two others, aged twenty
months and four years respectively, there was extensive tuberculous
enteritis with numerous ulcers and caseated mesenteric glands, and
in the second marked tuberculous peritonitis also; in each of these
there was also a lung focus. In the fifth, a girl of four years, there
was caries of three vertebrae, and several tuberculous ulcers were
present in the intestine; in the lower lobe of the right lung was a
focus as large as a cobnut.
It is a difficult matter to discuss this question apart from the
next, namely: —
5. The almost constant absence of evidence that the portal of entry
might be elsewhere when a lung focus was present .
In the previous points, my findings have been to a great extent
in agreement, but in this, if I understand it correctly, my experience
is distinctly at variance.
Several of the cases reported in this series would find an
explanation in a dual route of infection, whereas the statement above
39 °
would appear to rule out such an occurrence, not merely as a more
or less simultaneous infection, but even after an interval. So it
would seem to put forward the claim that, given a primary lung
focus due to respiratory route infection, there is little, if any,
likelihood of the intestine becoming affected unless secondarily to
the pulmonary focus. . This cannot be ascribed to an increased
resistance or immunity owing to the presence of the lung
tuberculosis, because it is a well-established fact that intestinal
tuberculosis can arise from the swallowing of infected sputum.
One explanation which, however, in my opinion, savours rather
of evading the difficulty than explaining it, would be to say that
whenever a lung focus is present together with definite alimentary
tuberculosis, the former was the primary site, and the intestinal,
though perhaps more advanced, arose from the swallowing of the
infected sputum. We may account for some of the cases in this
way: the respiratory route gives rise to the lung focus primarily,
the intestine becomes infected secondarily, and the lungs are again
involved by miliary tubercles spreading via the lymphatics to the
thoracic duct, and so to the pulmonary circulation. A considerable
number in this series may be thus explained, and several have been
mentioned in the previous paper.
In connection, however, with these points—the 1 almost constant
absence of a lung focus when the portal of entry app^urs to be
elsewhere,’ and the corollary of this, the 1 almost constant* absence
of evidence that the portal of entry might be elsewhere when a lung
focus was present ’—there are twelve which did not appear to
conform. There is no need to describe them all; two examples will
suffice: —(i) A girl of four years with tuberculous caries of the
spine. In the discussion on this case, it was stated: ‘ The spinal
site was probably the oldest; from the number of ulcers in the
intestine, from the fact of the large and the small both being
involved, jnd from the mesenteric glands being in large caseous
masses, the alimentary tract would appear to have been involved
* prior to the lung.* There was a sub-apical focus in the right lung
the size of a cobnut, becoming caseous, (ii) A female infant of
eight months; the mesenteric glands were in large caseous masses,
whereas the mediastinal were not much enlarged and contained
merely small caseating points. The tuberculous infection of the lung
39 1
was limited to the middle lobe of the right, in which there was a
distinct focus the size of a pea.
It has been stated by MacCallum that in children one may find,
instead of the apical lesion so common in adults, a caseous softening
of bronchial lymphatic glands and erosion through a bronchus to
produce wholesale tuberculosis of a lung, or a large section of it.
Several of the cases in this series might be looked upon as examples
of this; we must bear in mind, however, that, though this may
explain generalised infection of part or even the whole of a lung,
we still leave unaccounted for the source whence the gland became
involved- In those instances in which we find a focal lesion, the
subsequent more generalised condition in that lung, or part of it,
may be ascribed to reinfection from the gland, which itself was due
to the primary focal lesion. In cases where no focal lesion is found
to which the mediastinal gland could be traceable, and especially
in cases where both lungs are attacked by miliary or grey tubercles,
the spread may have occurred by the blood-stream; if general, % by
the pulmonary circulation, if localised, either by a branch of this
or of a bronchial artery. In one case, in which there was caries of
the right side of the sixth cervical vertebra and pus discharging
into the right pleural cavity, into a loculus shut off by pleural
adhesions, the glands at the hilus and along the trachea were
caseous and may have arisen secondarily to the lesion above, and the
widespread miliary affection of the right lung may then have resulted
by the method which MacCallum describes.
Passing on to the question of abdominal and alimentary tuber¬
culosis, there is not much to be said in this paper dealing with the
aspect of the morbid anatomy. As already stated, cases of isolated
primary tuberculosis of the intestines, not uncommon at home, are
comparatively rarely met with out here. In only four instances was
the disease confined to the abdomen; three of these were infants,
aged respectively ten, eighteen and twenty-two months, the fourth
was a child of seven years. From the intestines with, or more often
without, any local lesion, the bacilli are arrested for a time in
the mesenteric glands and thence spread either by way of the
lymphatics to the blood and so to the lungs (of this, several instances
have occurred in this series, and have already received sufficient
mention), or else by the portal blood to the liver. This has been
39 2
very rare in these cases. In fact, cases in which the liver was
extensively involved have been few. The spleen in nearly all was
more affected than the liver, and in two only was the liver infected
and not the spleen. When tubercles were found in the liver,
in the large majority of instances they were small, miliary to
pin-head, and confined to the surface. The rarer forms of intestinal
tuberculosis where the disease is localised to the region of the ileo-
caecal valve to produce a mass of tuberculous cicatricial tissue
constricting the lumen, I have met with onde among these cases.
The following is worthy of mention while dealing with alimentary
forms. The subject was a child of two years; there were tuberculous
ulcers in the small intestine, and the mesenteric glands were enlarged
and caseated. A tuberculous meningitis was the only other lesion
detected; there were numerous pin-head tubercles at the base and a
few on the vertex. The mode of extension in this case is obscure.
One only evades the question by saying that it was probably
metastatic by way of the blood-stream, analogous possibly to cerebral
abscess in cases of haemorrhoids ancj in liver conditions.
The meninges and brain were found involved in a considerable
number of cases, particularly the former, and usually as miliary
tuberculosis affecting mostly the base and the Sylvian fissures.
Definite focal masses in the brain or cerebellum were comparatively
rare.
There were several cases in which the lungs and meninges were
the only parts in which tubercles were found. In seven instances
the meninges and one lung only were affected; in six of these it was
the right lung which was attacked. In one other case, in addition
tq the lung and meningeal involvement, there was a tuberculous
focus the size of a small marble in the right cerebellar hemisphere.
In one case, a girl of eight years, the mediastinal glands and the
meninges were the only parts found affected, there being no tubercles,
either focal or miliary, in the lungs.
The mode of extension to produce the* peculiarly limited
distribution of tubercles in lungs and meninges is obscure, and is,
perhaps, analogous to brain affectioAs secondary to other pulmonary
conditions, bronchiectasis, for example. There is not, as far as
I am aware, any lymphatic connection between the lungs and the
base of the brain, and if the extension ( occurred by way of the
393
blood-stream, why should the secondary infection be so limited?
One would almost certainly expect to find signs in other organs.
I am also unable to suggest any significance for the fact that,
in the combination of lungs and meninges, of the seven cases in
which one lung only was affected, in six it was the right; and of
the five cases in which both were involved, in three the right was
more affected than the left. Again, in the case already referred to
in which the mediastinal glands and meninges were tuberculous,
but in which no signs were detected in the lungs, the glands were
those of the right side only.
The combination of lungs, meninges and spleen was a little more
common, fifteen of such being met with. In two of them only one
lung was involved, in each case the right.
In this connection, it may be worth noting that the involvement
of the spleen with miliary tubercles as the only abdominal viscus
affected has not been very infrequent, and may find explanation,
perhaps, if one regards the spleen as the meeting point of lymphatic
and blood terminals. I may refer here to the observations of Dumas
upon what he calls an unusual form of tuberculosis met with in
Salonica amongst Senegalese and Arab troops. He noted that the
mediastinal and peribronchial lymphatic systems were first affected,
and later the pleura and pericardium. In the early stages the
glands were merely enlarged, but later they suppurated or became
caseous. At the onset there were no lesions of the parenchyma of
the lung, and even later not the ulcerating and caseous form, only
a few scattered tubercles. He considered that it was in the spleen
that the lesions passed from the lymph to the blood stream. When
the spleen was normal no tubercles were found in the lungs, but
when the former was invaded the latter were also affected.
Examples of this in children, i.e., cases in which the spleen is
the only abdominal viscus involved and the connection referred to
between this and the pulmonary findings seem to find support in
some of the instances in this series. One may be given: a male
child, three years of age, showed miliary tubercles scattered
through both lungs, the spleen contained similar tubercles, as did
also the meninges. The hilus and paratracheal glands on the right
side were enlarged to the size of a cobnut and were caseous. Again,
as was pointed out above, in the peculiar limitation at times to the
m
lungs and meninges, if haematogenous metastasis occurs, the first
part to suffer is the meninges. In two other instances, however,
the lungs were affected with small tubercles (no focus) and the hilus
and paratracheal glands were caseated, but the spleen was not
involved; in one of these blood infection was apparent, since the
meninges revealed basal tubercles.
On nine occasions were focal tubercles found in the brain or
cerebellum. The commonest site was the latter, for cerebellar were
found in eight. Of these, the focus was present in four cases in tire
left hemisphere, in three in the right, and once in both. In this
last there were four distinct foci, two in each lateral lobe; two others
had a second, extra cerebellar, focus, namely, in the right cerebral
hemisphere in one, in the left crus in the other. Finally, one
subject, a female child of two and a half years, exhibited multiple
foci: three distinct ones in the left lobe of the cerebellum, two in the
left cerebral hemisphere, and one in the left hippocampus.
A striking feature of the tuberculous conditions as met with here
is the rarity of bone, joint or skin affections. In this series there
were five with tuberculous ulcers of the skin, four of these on the
face and neck and one on the forearm. Not a single instance of
tuberculous joint disease was encountered, and only three with bone
lesions. In two of these there were caries of the lower dorsal and
upper lumber vertebrae; a third had caries of the sixth cervical
vertebra, and tuberculous disease also of the left tibia and both
femora. A fourth had widespread tuberculous disease—lungs,
liver, spleen, peritoneum and meninges—and extensive caries of the
left mastoid, but there was no absolute proof that this last was due
to tubercle.
Tuberculous cervical adenitis, so frequently met with at home,
is a comparative rarity in the post-mortem room here. Of the whole
series there were only twenty-eight instances, and in three of these
the enlargement was very slight, so slight that there was not
sufficient macroscopical evidence to determine its nature; on section,
however, giant cells and tubercle bacilli were seen.
In one case there was almost universal involvement of the
glands—cervical, submaxillary, supraclavicular, axillary, media¬
stinal, mesenteric and femoral—together with widespread pulmonary,
intestinal, and some renal tuberculosis. This was a child of only
395
three years. Sections did not show any of the usual conditions
characteristic of Hodgkin’s disease. In contrast to this, may be
incidentally mentioned a little girl of seven months, showing
but one cervical gland enlarged, with very extensive tuberculous
disease—lungs, pleura, mediastinal glands, mesentery, intestines,
liver, spleen, kidneys, meninges.
Cases in which the genito-urinary tract has been involved have
been comparatively few in my experience here. The majority of
these showed a few miliary tubercles as emboli in the glomerular
capillaries, consituting part of a general haemic infection. There
were ten instances in which the kidneys were found foe ally affected
in conjunction with widespread tuberculosis. One, a girl of
three years of age, merits further mention on account of the
peculiar distribution. The extensive alimentary infection—tuber¬
culous ulceration of both large and small intestines, caseated,
adherent mesenteric glands, extensive infection of the peritoneum—-
and the equal distribution of tubercles throughout both lungs would
point to the alimentary canal as the primary portal of entry. The
condition of the kidneys, however, does not support the idea of a
haematogenous origin from entrance of the bacilli into the general
circulation from the lungs. Each kidney showed a large focus in
a similar situation at the lower pole, the size of a cobnut and
caseous, most advanced at the margin between the cortex and a
pyramid, and passing in as if later it would discharge into the
pelvis. No indication of involvement of the ureters was detected.
The foci gave the impression that they had arisen from affection of
the tubules in course of excretion (as mentioned by Aschoff and
Israel). The foci were each of them, in a more advanced state than
a few smaller, pin-head and miliary, tubercles in the cortical area
of the right kidney, which had more likely arisen by haematogenous
infection. The kidney foci appeared to be of considerably older
standing than the pulmonary condition, and showed more advanced
caseation than the mediastinal glands, but less than the mesenteric.
There were no indications of an ascending infection from lower
down the urinary tract. Other viscera, liver and spleen, showed
sparse tubercles only, and these were, in the former at all events,
confined to the peritoneal surface. Briefly, the age of the kidney
condition appeared to be less than that of the alimentary but
396
more than that of the pulmonary, and the mode of involvement is
obscure.
Two others are of sufficient interest to warrant brief mention: —
(i) An infant of nine months, showing generalised tuberculosis-
lungs, intestines, liver, spleen, kidney—and, in addition to miliary
infection of the last, one calyx in the left was hollowed out and
lined by tuberculous material; (ii) a girl of four years, with miliary
tuberculosis of the lungs and of the liver surface, but the only focus
found in the body was a caseated mass occupying practically the
whole of a pyramid in the left kidney.
Finally, while speaking on the subject of gemto-urinary
affection, mention must be made of two cases of exceptional interest.
Both exhibited extensive disease, lungs, meninges, intestines,
peritoneum and ovaries. In one, a girl of four years, both ovaries
were enlarged, and had become converted practically into caseated
masses; this case has been described in a previous paper when
discussing the primary portal of entry. In the other, an infant of
only eight months, the Fallopian tubes were swollen and caseous on
both sides, while the ovaries were in a condition similar to the last.
In this case a possible (or, rather, probable) source was by contiguity
from the tuberculous peritoneum, but in the former the tubes did
not appear to be affected.
REFERENCES
Aschoit and Israel (1919). Cited by MacCallum. Text-book of Pathology, p. 616.
Bushnill, G. E. (1918). Military Surgeon .
Calmette, Guerin, and Briton (1907). Ann. it VInstitut Pasteur , Vol. XXI, No. 6.
Canti, R. G. (1919). Primary Pulmonary Tuberculosis in Children. Quarterly Joum. Med-
Vol. XIII, No. 49, pp. 71-81.
Delepine, S. Contribution to the Study of Delayed or ‘latent' Tuberculous Infection.
Ann. it rinstttut Pasteur .
Dumas, A. (1919). Adenopathie trach^o-bronchique A marche rapide. Lyon Med ., pp.
180-187.
Ghon, A. Die primare Lungenherd bei der Tuberkulose der Kinder.
MacCullum, W. G. (1919). Text-book of Pathology , p. 607.
397
ON THE GENUS CYLICOSTOMUM*
BY
J. E. W. IHLE
(From the Zoological Laboratory , Veterinary College ,
Utrecht)
(Received for publication 29 October , 1921)
Last year (1920 a, p. 268) I published an enumeration of the
species of the genus Cylicostomum, and divided this genus into,
eight groups, of which three had been distinguished by Looss
(1902, pp. 130-132). Since that time some new species have been
described. Moreover, I think it necessary to modify somewhat the
groups, so that I now divide the Cylicostomum species as follows: —
I. Tetracanthum-coronatum group.
The external leaf-crown is composed of eighteen to twenty-four
elements; those of the internal leaf-crown are thin, triangular plates,
the place of origin of the latter extending in a backward direction
to some distance from the anterior margin of the mouth-capsule.
Mouth-capsule rather short. Posterior extremity of the $ straight
or slightly bent in a dorsal direction.
• Of late, tome authors (Railliet, Travassos) use, instead of the name Cylicostomum , the
genus-name Tricbonema , which ought to be used according to the law of priority, In 1874,
Cobbold (Veterinarian, Vol. XLVII, p. 85) described under the name Tricbonema arcuata
larvae of Cylicostomum. One year later ( Veterinarian , Vol. XLVIII, p. 241) he established
the fact that this Tricbonema is the larva of ‘ Strongylus tetracantbus the present genus
Cylicostomum Railliet. As, however, the law of priority must be applied * when any stage in
the life-history is named before the adult,' and the name Cylicostomum (Railliet, L ' £cbo Viterin M
Vol. XXXI, p. 40) is used in 1901 for the first time, Tricbonema ought to be used.
It does not seem advisable to me in this, nor in the like cases, to adhere to the rules of
zoological nomenclature, when, owing to this, a completely familiar name would be replaced
by one still entirely unknown. Neither did I always adhere to the law of priority in my revision
of the Trematodes, Cestodes and Nematodes ot domestic animals, in the third edition of Sluiter
and Swellengrebel, 1 De dierlijke parasieten van*den mensch en van onze huisdieren.*
39 ®
1. C. tetracanthum (Looss), 1902, p. 124.
2. C. labratum (Looss), 1902, p. 124. H.*
3. C. ornatum, Kotl&n, 1919, p. 10.
4. C. labiatum (Looss), 1902, p. 125.
4 a. C. labiatum (Looss) var. digitatum, n. var. H.
5. C. sagittatum, Kotlin, 1920, p. 4.
6. C. corona turn (Looss), 1902, p. 125. H.
II. Alveatum-catinatum group.
The external leaf-crown consists of twenty to twenty-nine
elements; those of the internal leaf-crown are similar to those of the
first group. Posterior extremity of the $ strongly bent in a dorsal
-direction, with a swelling before the vulva, so that it reminds one of
a human foot, seen laterally. .
7. C. alveatum (Looss), 1902, p. 127.
8. C. catinatum (Looss), 1902, p. 128.
8 a. C. catinatum (Looss), var. litoraureum, Yorke and Macfie,
1920, p. 166.
9. C. pseudocatinatum, Yorke and Macfie, 1919, p. 273. H.
10. C. pat era turn, Yorke and Macfie, 1919 a, p. 57 (= C. cyma-
tostomum y Kotlin, 1919, p. 11). H.
11. C. goldi t Boulenger, 1917, p. 210. H.
12. C. tridentatum , Yorke and Macfie, 1920, p. 153.
13. C. mettamiy Leiper, 1913, p. 460.
C. tridentatum and C. goldi are evidently closely allied
species, as I found the three teeth of the oesophageal funnel of
C. tridentatum , described by Yorke and Macfie, also in C. goldi.
They are visible only in thoroughly transparent specimens with dean
mouth-capsule.
It seems to me that C. pseudocatinatum must be considered a
variety of C. catinatum. The former does not differ more from
C. catinatum than does C. catinatum var. litoraureum , Yorke and
Macfie. Both differ from C. catinatum in the appendages of the
genital cone. Moreover, the appendages of C. catinatum litoraureum
are more like those of C. pseudocatinatum than like those of the
typical catinatum form. So when we consider litoraureum a variety
as Yorke and Macfie do, this also applies to pseudocatinatum .
The ipecie# occurring in Holland arc marked H.
399
III. Radiatum-elongatum group.
Mouth-capsule with a hoop-like thickening at the posterior
margin. Elements of the internal leaf-crown generally small and
numerous, originating close to the anterior margin of the mouth-
capsule. Posterior extremity of the 9 straight, or slightly bent in
a dorsal direction.
14. C. radiatum (Looss), 1902, p. 129.
15. C. triramosum, Yorke and Macfie, 1920, p. 175.
16. C. elongatum (Looss), 1902, p. 129. H.
16 a. C. elongatum (Looss) var. Kotlani, Ihle, 1920, p. 269
(= C. elongatum var. macrobursatum, Kotldn, 1920,
p. 6). H.
17. C. insigne, Boulenger, 1917, p. 207. H.
18. C- tebtae, Boulenger, 1920 a, p. 102.
19. C. adersi, Boulenger, 1920, p. 30.
20. C. nassatum (Looss), 1902, p. 128.
20a. C. nassatum (Looss) var. parvum, Yorke and Macfie
(1918, p. 400). H.
21. C. leptostomum, Kotlin, 1920, p. 3. H.
22. C. auriculatum (Looss), 1902, p. 130.
The 9 of the species last mentioned forms a transition to the
alveatum-catiuatum group by the form of its posterior extremity.
In this group I now also include C. leptostomum, Kotldn, a
species about which Kotldn published a description in Hungarian
only; the author was so kind, however, as to send me a German
translation. C. leptostomum is closely allied to C. nassatum, from
which it is distinguished by the posterior extremity of the 9 being
straight.
IV. Calicatum group.
Mouth-capsule mostly cylindrical or trapezium-shaped in optical
section. Elements of the internal leaf-crown mostly short and
implanted in the immediate neighbourhood of the anterior margin
of the mouth-capsule. Posterior end of the 9 mostly straight.
23. C. calicatum (Looss), 1902, p. 127. H.
24. C. minutum, Yorke and Macfie, 1918, p. 405 (= C. cali¬
catum var. minus, Kotldn, 1920, p. 6). H.
400
25. C. longibursalum , Yorke and Macfie, 1918, p. 400
(= C. calicatiforme , Kotl&n, 1919 a, p. 559 = C. nanum,
Ihle, 1919, p. 720). H.
26. C. hybridum , Kotlan, 1920, p. 5.
27. C. poculatum (Looss), 1902, p. 126. H.
The last mentioned species differs considerably from C. calicatum
( e.g. t in the bursa, which is closed all round and has a finely
denticulated border) so that Looss (1902, p. 132) did not group
them under one head.
V. Euproctus-bicoronatum group.
The external leaf-crown mostly consists of numerous slender
elements; the elements of the internal leaf-crown are extremely
large. The posterior extremity of the 9 is straight.
28. C . euproctus , Boulenger, 1917, p. 204. H.
29. C. bicoronatutn (Looss), 1902, p. 125. H.
30. C. ihlei , Kotlan, 1921, p. 300. H.
31. C . ultrajectinum , Ihle, 1920a, p. 269; 1921, p. 372. H.
For the present, some heterogenous elements are taken together
in this group, characterized by the possession of very large elements
of the internal leaf-crown.
C. ihlei is closely allied to the genus Poteriostomum in structure
of mouth-capsule and leaf-crowns. The fact, however, that the
bursa copulatrix of this species shows the typical characteristics of
Cylicostomum and differs from that of Poteriostomum , supports the
opinion, shared by Yorke and Macfie (1920, p. 159), that Poterio¬
stomum must be considered an independent genus contrary to
Kotl&n’s view (1921, p. 299).
Formerly, I considered C . ultrajectinum to belong to the
radiatum-elongatum group, with which it agrees in the thickened
posterior margin of the mouth-capsule, from which it differs,
however, in the size of the elements of the internal leaf-crown. It
is, however, also considerably different from the other species erf the
euproctus-bicoronatum group in size and number of the elements
of the external leaf-crown. Yorke and Macfie (1920, p. 162)
expressed the opinion that C. ultrajectinum might belong to the genus
4oi
Poleriostomum . It has become evident, however (Ihle, 1921,
p. 372), that the bursa copulatrix resembles that of the other
Cylicostomum species in every respect, so that it is certain that this
species belongs to the genus Cylicostomum .
VI. Brevicapsulatum group.
Mouth-capsule extremely short. Posterior extremity of the 9
straight.
32. C . brevicapsulatum , Ihle, 1920, p. 562. H.
33. C . prionodes , Kotldn, 1921, p. 305.
These two species differ considerably, but they can be grouped
together for the present.
VII. Montgomeryi group.
The dorsal and ventral walls of the mouth-capsule are much
longer than the lateral walls.
34. C . montgomeryi , Boulegger, 19200, p. 104.
I will conclude by describing a new variety of C . labiatum
(Looss), of which I found a small number of specimens in the large
intestine of the horse in Holland.
Cylicostomum labiatum (Looss) var. digitatum , nov. var.
The new variety agrees in the main (mouth-collar, mouth-
capsule) with the typical form.
<3 7 to 8 mm., the immature 9 8 to 8f mm. long. Oesophagus
345 to 410ft long. The bursa copulatrix has a dorsal lobe, which
varies in length and is mostly longer than in the typical form. The
distance from the extremity of D 3 * to the point of origin of the
postero-extemal ray varies from 310ft to 380/1. The posterior part
of the dermal collar shows the processes occurring in the typical
form. The chief difference from the typical form is that the new
variety shows finger-like processes on both sides of the posterior
part of the genital cone, which are absent in the typical form. The
number and form of these processes perhaps fluctuate. (In one case
I counted three on each side.) The posterior extremity of the 9
is like that of the typical form. The distance from the vulva to the
• The short stem which the dorsal rays have in common, I name D®, the two main trunks D,
the side branches, from before backwards, D*, D*, and D*. In Cylicostomum D s forms the
immediate continuation of D,
402
anus is smaller however (ca. 95 a 1 ) than in the typical form; the
distance from the anus to the posterior extremity is 86/1 to I io ft,
Fig. i. Cylicostomum labiatum Fig. 2. Cylicostomumlabiatum Fig. 3. CyHcosfmsm lakiatm
var. digitatum. Dorsal lobe of the Far. digitatum. Processes of the var. digitatum Posterior extremity
bursa. genital cone, lateral view. of female, lateral riew.
measured along the axis of the body. The conical end of the body
is bent somewhat in a ventral direction.
REFERENCES
Boulenger, C. L. (1917)- Sclerostome parasites of the horse in England. II : New species
of the genus Cylicbnostomum. Parasitology. Vol. IX.
-(!92o). Sclerostomes of the donkey in Zanzibar and East Africa. Parasitology ,
Vol. XII.
- (1920 a). On some Nematode parasites of the Zebra. Parasitology , Vol. XII.
Ihle, J. E. W. (1919). Cylicostomum nanum , een nieuwe Strongylide van het paard. Tijd-
scbrijt voor Diergeneeskunde. Dl. 46.
- (1920). Cylicostomum brevicapsulatum , n.sp., eine neue Strongylide aus dem Darm
des Pferdes. Ccntralblatt f. Baht. Abt. 1. Orig. Bd. LXXXTV.
- (1920 a). Bemerkungen ubcr die Gattungen Cylicostomum , Poteriostomum und
Craterostomum. Centralbl.f. Bakt. Abt. 1. Orig. Bd. LXXXV.
- (1921). Das Mannchen von Cylicostomum ultrajectinum. Centralbl.f. Bakt. Abt. 1.
Orig. Bd. LXXXV.
KotlAn, A. (1919). Beitrag zur Helminthologie Ungarns. I. Neue Sclerostomiden aus
dem Pferd. Centralbl.f. Bakt. Abt. 1. Orig. Bd. LXXXIII.
- (1920). (Hungarian paper). Allatorvosi Lapok 1920.
- (1921). Two new Cylicostomum species of the horse. Am. Trap. Med. & Par ant.,
Vol. XIV.
Leiper, R. T. (1913). A new Cylicostome worm from the horse in London. Veteriu. Jouns.,
Vol. LXIX.
Loom, A. (1902). The Sclerostomidae of horses and donkeys in Egypt. Bee. Egypt. Govern.
School of Med., 1901.
York*, W., and Macfie, J. W. S. (1918). Strongylidae in horses. I. Cylicostomum longi-
butsatum , sp.n. ; II. C. minutum , sp.n. ; III. C. a ass a turn Loots var. parvusm. Ann Trop.
Med. & Parasit ., Vol. XI.
- (1919). Idem. VI. C. pseudo-catinatum , sp.n. Ibid , Vol. XII.
- (1919 a). Idem. VII. C. pateratum , sp.n. Ibid, Vol. XIII.
-(* 9 2 °)* Idem. IX. Cylicostomum tridentatum , sp.n.; X. On the genus
Poteriostomum (Quiel). XI. Species found in West Africa and Jamaica; XIII.
Cylicostomum triramosum , sp.n. Ibid\ Vol. XIV.
4°3
NOTES ON AUSTRALIAN CESTODES
BY
P. A. MAPLESTONE
{Received for publication 17 November, 1921)
INTRODUCTION
Whilst on the staff of the Australian Institute of Tropical
Medicine, Townsville, a systematic examination was made of the
large collection of Cestodes in the Museum. In addition to a
considerable number of new species, the description of which will
appear in subsequent papers, a number of previously described
worms were found in new hosts.
I. PREVIOUSLY DESCRIBED CESTODES IN NEW HOSTS
Cittotaenia tackyglossi, Johnston, 1911
= Cittotaenia , sp. nov., Nicoll, 1914.
Nicoll (1914) recorded a new species of Cittotaenia from the
Echidna, Tackyglossus aculeatus, Shaw, but without furnishing any
description of it. Detailed re-examination of the same material
proves it to be Cittotaenia tacky glossi, Johnston, but the dimensions
are all somewhat greater than Johnston gives in his description,
notwithstanding the fact that the worms are in an immature state
similar to those from which the original description is given.
Diorchis flavescens (Krefft), Johnston, 1912.
The original description of this worm was published by Krefft
(1873), who named it Taenia flavescens, but H. Johnston (1912) 1
re-examined Krefft’s material and placed it in the genus Diorchis.
Up to the present, this cestode has been found in Anas superciliosa,
Gmel. r the Black Duck; Spatula rhynchotis, Lath., the Blue-winged
Shoveller; Nettion castaneum, Eyton, the Teal; and Aythya
+°4
australis, Gould, the White-eyed Duck. A few badly preserved
fragments of apparently the same species have been found by the
writer in material taken from Dendryocygna arcuata, Cuvier, the
Whistling Duck.
Ophiotaenia longmani, Johnston, 1916.
Johnston (1916) originally described this species from the
snake Aspidotes ramsayi , which was. taken at Yarelba, Western
Queensland. The same species was recently found by the writer in
the intestine of Python spilotes var. variegatus, Gray, the Carpet
Snake, which was killed near Townsville, North Queensland.
Ophiotaenia hylae, Johnston, 1912.
O. hylae was described by Johnston (1912) 2 with the frog, Hyla
aurea, as its host, which was captured near Sydney, New South
Wales. A scolex and several proglottides of this species were found
in the intestine of D. arcuata , the Whistling Duck, which was shot
near Townsville. This is strange, as the family Proteocephalidae ,
to which the above species belongs, has hitherto been recorded only
in Amphibia, Reptilia, and the dog. The most probable explana¬
tion of this occurrence is, that the duck had swallowed a frog
infected with the cestode, and the frog had been sufficiently digested
to liberate it into the bird’s intestine, without itself having undergone
digestive changes up to the time the duck was shot.
Acanthotaenia gallardi , Johnston, 1911.
This cestode was originally described by Johnston (1911) 2
under the name Proteocephalus gallardi, and subsequently placed
in the genus Acanthotaenia. On this occasion it was found in the
intestine of Pseudechis porphyriacus, Shaw, the Black Snake.
Since then it has been recorded by Johnston (1912) 3 as occurring
in Pseudechis australis, Gray, the Northern Black Snake; Notechis
scutatus, Peters, the Tiger Snake; and Denisonia superba* Gunth,
the Copper-headed Snake. To this list of hosts the writer is now
able to. add Dipsadomorphus fuscus’ Gray, the Brown Tree Snake,
4°S
a specimen of this snake harbouring this worm being killed near
Townsville.
Moniezia alba, Blanchard, 1891.
Johnston has recorded the presence of this cestode in Australia
in sheep in New South Wales, and the writer has recently obtained
specimens of the same worm from a bullock, slaughtered in
Townsville, and which came from near Hughenden, Western
Queensland. This is the first record of this cestode in the above
host in Australia.
Thysanosoma giardi , Stiles, 1893.
Johnston has noted the occurrence of this worm in sheep in
New South Wales, but no note of its frequency and distribu¬
tion has been given. Over a period of several months, the
writer has had the opportunity of examining many worms taken
from sheep in Townsville, and they all proved to be of this species.
The sheep came from districts representing a wide area in Western
Queensland, so it seems that it is widely distributed, and very
common, but as far as could be ascertained from inquiries among
pastoralists and butchers, it does not give rise to the serious patho¬
logical condition among sheep, that it does in other parts of the
world.
REFERENCES
Johnston, T. Harvey (1911).! Ccstoda and Acanthocephala. Report Aust. Inst. Trop. Med.
- (1911).* A new Cestode from the Black Snake. Annals of the Queensland Museum,
No. 10, p. 1.
- (1912)4 Re-examination of the types of Krefft's species of Cestodes. Rec. Aust.
Museum, Vol. IX, No. i.
- (1912).* A Census of Australian Reptilian Entozoa. Proc. Roy. Soc., Queensland, Vol.
XXIII, p. 233.
- (1912).* Notes on some Entozoa. Proc. Roy. Soc., Queensland, Vol. XXIV, p. 63.
- (1916). Helminthological notes. M. Queensland Museum , Vol. V, p. 186.
Kjofft (1873). Trans. Entomological Soc. N.S. Wales, II, pp. 206-227.
Nicoll, W. (1914). Remarks on the worm Parasites of Tropical Queensland. Med. Journ.
of Aust., p. 244.
4°7
NOTES ON AUSTRALIAN CESTODES
BY
P. A. MAPLESTONE
(Received tor publication 25 'November , 1921)
II. ANGULAR!A AUSTRALIS , sp. nor.
*
This cestode was found in considerable numbers in the intestine
of a Stone Curlew (Burhinus graUarius , Lath.), shot near Townsville,
North Queensland. It is probably rather rare, this being the only
occasion on which it was encountered.
External Anatomy.
The worms are very small; the largest individual measured only
about 3 mm. long; from its appearance the strobila was complete,
but it had not quite reached full development. The number of
proglottides in a chain varies between fourteen and twenty (fig. 1).
Fig. 1. A. australis. Appearance of worm at a whole. X 35.
On account of its small size, there are no macroscopic characters by
which it may be recognised, so the following description of the
appearance as a whole had to be determined under the low power of
the microscope.
408
Head . The head bears a relatively long, thin rostellum, which
was unfortunately broken off in most cases. From base to tip the
rostellum is about 150/1 to 200/1 long, and it is about 14/1 thick. It
arises from a bulbous muscular structure, which lies in a deep fossa.
It projects directly forwards as a long proboscis-like organ of uniform
diameter, except at the tip where it expands into a globular enlarge¬
ment, about twice the diameter of the stem (figs. 2 and 3).
Fio. 2. A . australis. Head, and Fig. 4. A. australis. Hooks, x 450.
anterior parts of strobila. x 70.
The hooks, which measure about 25/1 in length, are arranged in
a zigzag line which forms four acute angles anteriorly and four acute
angles posteriorly. 4 Between an anterior and a posterior angle
there are six or seven hooks. Thus, the total number of hooks is
forty-eight or fifty-six. Their disposition and shape are shown in
figs. 3 and 4.
409
The scolex is about as broad as it is long (250 /i). The four
suckers are relatively large oval structures, with their long axes
antero-posterior; they are not exactly oval, being slightly narrower
anteriorly. They measure about 130/1 in length and 60 /a in breadth,
and their lips are provided with cuticular expansions, which project
beyond the surface of the scolex (fig. 2).
Segments . For about the anterior half of the strobila the
segments are quite narrow and rudimentary, the only change being
a slight increase in length. But from about the mid-point of
the strobila, the proglottides rapidly increase in size, and the
reproductive organs reach complete maturity in the course of three
or four segments. There are only three or-four mature segments in
each strobila, the genitalia undergoing a sudden atrophy when the
uterus begins to develop, so that the two or three terminal segments
contain, beside the uterus, only remnants of the reproductive organs
(fig- 0-
The shape of individual segments is somewhat uncommon.
Each one is like a truncated cone with the narrow end anterior, and
the sides, which are concave in the anterior immature proglottides,
become slightly convex in the posterior mature and gravid segments.
The posterior surface of each proglottis is oval and slightly
depressed in the centre, and into the middle of this depression the
narrow anterior margin of the succeeding segment fits; the result of
this is that the posterior borders project all round beyond the
anterior portion of the following proglottis. There is no neck, but
posteriorly the scolex narrows considerably to pass directly into the
segmented chain. The first two or three proglottides are distinctly
broader than long, and the next three or four increase in length
but not in breadth, so that the sixth or seventh segment is longer
than broad. Mature and gravid segments are slightly broader
than long, their dimensions are about 160/1 antero-posteriorly,
270/1 across the anterior, and 460^ across the posterior borders
respectively. But these dimensions are only approximate, on account
of the rapid development of the worm. The terminal segment is
nearly globular, with an invaginated pore at its posterior end.
410
Internal Anatomy.
Muscular system . The muscle layers are not conspicuous, and
as sections were not cut, their detailed arrangement cannot be given.
Nervous system. The nervous system was not investigated.
Excretory system. The excretory canals, of which only a single
one could be seen on each side, lie well towards the lateral borders,
and pass ventral to the cirrus pouch and vagina.
Genitalia. Testes. The testes are arranged in the two lateral
fields of the medulla with the female organs between them. They
number about twenty in each segment, and there are usually one or
two testes more on the aporal than on the poral side. They lie
towards the posterior part of the segment on each side of the ovary
(fig. 5). The cirrus pouch is relatively long and runs slightly
4.M. e^dAi.
Fig. 5. A . australis. Mature segments, e.v., excretory vessel; c.p., cirrus pouch;
c., cirrus; t., teste#; v.d., vas deferens j r.s., receptaculum seminis ; o., ovary; v.g., vitelline
gland; v., vagina. X 90.
anteriorly towards the median line.’ The genital pores are situated
about the centre of the lateral border, and are regularly alternating
in most cases, but in one or two specimens there were three openings
in succession on the same side, which was the greatest irregularity
observed. There is no external seminal vesicle, but the vas deferens
4 11
is thrown into many coils in front of the ovary,* before it enters the
base of the cirrus pouch (fig. 5).
The cirrus is a long tubular structure, and in the one instance
observed, where it was extruded, it measured 120 p long and
i&P thick. It is thickly covered with spines, and has a slightly
swollen tip.
Ovary . The ovary lies in the median axis towards the posterior
part of the proglottides. It is a compact, oval body; its long axis
is transverse, measuring about 75/1 (fig. 5).
Vitelline glands . The vitellarium is small and lies transversely
behind the ovary, at the extreme posterior part of the segment; in
full development it measures about 40/1 long (fig. 5). The ovary
and vitelline glands disappear very suddenly, being apparently in
full development in one segment, and almost totally absent in the
succeeding one.
Vagina . The vagina is a narrow tube running posterior to
the cirrus pouch. It expands in front of the ovary into a small
receptaculum seminis. Details of the shell gland could not be
made out.
Uterus and eggs . The uterus was not fully developed, and its
mature characters cannot be detailed, but it had the appearance
of an irregular sac loosely packed with eggs, and is only visible
in at most the three terminal segments. No free eggs were seen,
consequently their size is not known.
Diagnosis.
This interesting cestode closely resembles Angularia beema,
Clerc (1906), but differs from the latter in the following points: —
(1) Its smaller size (A. beema measures 45 mm. long), and (2) the
presence of cuticular expansions on the suckers (absent in A. beema).
It is, therefore, proposed to name this species Angularia australis.
The discovery of this cestode is of considerable interest, because
the only other member of this genus hitherto described is A. beema ,
which, according to Liihe (1910), is very restricted in its distribution,
only having been recorded from Russia.
Type specimens of this new species are in the Museum of the
Liverpool School of Tropical Medicine.
The genus Angularia , Clerc (1906), resembles closely the genus
4 12
Gyrocoelia , Fiihrmann (1899), so far as the head is concerned.
Apparently, Linstow's genus Brochocephalus (1906) is a synonym oi
Fiihrmann’s Gyrocoelia . The points in which the two genera differ
may be summarised as follows : —
A ngularia.
Vagina present.
Testes 20 to 25.
Uterus without dorsal and
ventral pores.
Gyrocoelia,
No vagina.
Testes few.
Uterus ring-like, opening by
a pore dorsally and ven-
trally in gravid segments.
REFERENCES
Clerc, W. (1906). Notes sur les cestodes d'oiseaux d'Oural. Centralbl. Baht ., Orig. Abt. x.
Vol. XLII, No. 5, Oct. 12, pp. 728-730. Jena.
Linstow von (1905). Helminths from the collection of the Colombo Museum. Spclia
zeyhmicO) Colombo.
L&he, M. (1910). Die Sfisswasserfauna Deutschlands. Cesloda. Heft. 18, p. 1x5. Jena.
4'3
NOTES ON ULCERATIVE GRANULOMA
BY
P. A. MAPLESTONE
(Liverpool School of Tropical Medicine)
(Received for publication 17 November , 1921)
The following notes on cases of Granuloma among Australian
aborigines may be of interest, as they present some rather unusual
features. The patients were treated at the Australian Institute of
Tropical Medicine, Townsville.
CASE i. Male, aet. about 50. The disease was first noticed
about two years before admission to hospital; it began on the usual
site, viz., the penis, and spread slowly and continuously. After this
lesion had existed for some months, an ulcer appeared under the
chin in the fold of the neck.
When first examined, the case presented the following
appearance. The ulcer, which began on the penis, had spread until
it reached from one anterior superior iliac spine to the other, and
the penis itself had sloughed completely away, leaving the urethra
to open in the middle of the ulcerated surface. The ulcer on the
neck was about one and a half inches wide, and extended across
under the chin, from one angle of the jaw to the other, the sub¬
maxillary lymphatic glands on both sides were considerably swollen,
with consequent difficulty in opening the mouth.
The two lesions were similar in appearance, having clear-cut
edges and shallow bases; they did not exhibit any granulomatous
masses on their surfaces. Diagnosis in both ulcers was established
by the discovery in them of the bodies described in Ulcerating
Granuloma by Aragao and Vianna (1913).
This case presented the uncommon condition of a secondary
focus of disease at a site distant from the primary, whither it had
been most likely carried by the patient’s hands.
CASE 2. Female, aet. 22. About three months prior to
4H
admission to hospital the woman noticed a small nodule on the
inner surface of the right upper lip; the nodule soon ulcerated, and
the swelling and ulceration spread fairly rapidly. When first seen,
there was a swelling of considerable size, which extended from the
right upper lip, over about the anterior half of the inner side of the
right cheek; there was more tumefaction and less ulceration than is
generally seen in genital granuloma of the same duration. Smears
made from scrapings of this growth were found to contain very
numerous organisms, similar to those found in Case I.
Typical infected large mononuclear cell from Case 2. X 1125.
There was no sign of a lesion in the genital region, so the case
is apparently one of primary infection in a most unusual position.
CASES 3 and 4. These were both young native women. They
both had small typical granulomatous lesions on the vulva, which
they stated had only been present for two or three weeks. On
examining the hospital records, it was found that both these women
had been treated for similar conditions previously, and that one
had been discharged fourteen months and the other fifteen months
previously, both being apparently cured. A further striking fact,
was that on the second occasion the lesions were on exactly the
same site as on the first occasion, in both cases.
These two may be cases of reinfection after cure, but the
similarity in time of the second manifestation of the disease, coupled
with the fact that the lesions are both in exactly the same place as
on the first occasion, makes it appear that they are recurrences after
apparent cure.
TREATMENT
The routine treatment adopted in all cases was the application
of a simple, non-irritant, antiseptic dressing to the lesions, and the
intravenous administration of tartar emetic; i grain in io c.c. of
normal saline twice weekly. Larger doses were unnecessary,
improvement being observed after the first or second injection, and
healing continued without signs of relapse during the course of
treatment. The time required for complete disappearance of the
lesions was proportionate to the extent of the ulceration present;
e.g. y the first case cited above, with two large ulcers, took about
three months to heal, whereas Case 2, with a small amount of
ulceration, healed in about a month.
On account of the tendency of the disease to recur, five or six
injections were given after the ulcer had quite healed.
REFERENCE
Aragao, H. de B., and Vianna, G. (1913). Peiquizaa *obre 0 Granuloma venereo. Mem. do
Instit. O ste al do Cruz, Vol. V. Rio dc Janeiro.
4'7
THE STRUCTURAL DIFFERENCES
IN THE OVA OF ANOPHELES
MACULIPENNIS , A. BIFURCATUS
AND A. PLUM BE US
BY
MALCOLM E. MacGREGOR
Wellcome Field Laboratory, Wisley, Surrey
{Received for publication \g November, 1921)
Plate XXII
CONTENTS
4*7
... 418
4*9
421
421
422
424
i. Introduction .
ii. Certain Characters or Mosquito Ova
iii. The Ova or Anopheles maculipennis
iv. The Ova or Anopheles bifurcatus .
v. The Ova or Anopheles plumheus ...
vi. Facts connected with Oviposition
1. Anopheles maculipennis.
2. Anopheles bifurcatus.
3. Anopheles plumheus.
vii. Reverences. .
INTRODUCTION
During investigations which entailed the breeding and rearing
of the three species of British Anophelinae at the War Office
Laboratory at Sandwich, Kent, in the summer of 1919, I noticed the
quite different appearances between the eggs of Anopheles maculi¬
pennis, A. bifurcatus , and A. plumheus, perceptible even to the
unaided eye.
There are numerous illustrations of Anopheline ova in the English
mosquito literature, but I have seen none indicating the markings on
the dorsal surface of the ova of A . maculipennis which make it so
easy a matter to identify the ova of this species from those of
A. bifurcatus .
4i8
CERTAIN CHARACTERS OF MOSQUITO OVA
The eggs of mosquitoes, although showing much difference in
form among the various species, nevertheless have certain characters
that are uniform. Generally they are elongated, roughly cigar¬
shaped, or oval objects. The living elements in the eggs of insects
are protected by three separate external coats: (i) a delicate
innermost coat termed the vitelline membrane; (2) the hard, shell-like
materia] termed the chorion; and (3) a thin outermost membrane
which I shall refer to as the enveloping membrane.
All mosquito ova have the egg-shell proper (chorion) covered or
partially covered with the delicate semi-transparent and water-proof
enveloping membrane.
This membrane envelopes the egg but is not closely adherent
to the shell, a layer of air often existing between the shell and
membrane.* Eggs may be easily freed from the membrane by means
of pressure with the point oTa blunt needle, or by boiling the eggs
in water for a moment or two. This membrane may either conform
to the surface of the egg and lie so close to the egg-shell as to be
only visible when special methods are adopted to demonstrate its
presence, or it may, by being * ballooned \ form structures around
the egg. Such structures are seen in the 1 floats 9 of Anopheline ova.
The eggs of mosquitoes may be divided into two classes: (1) those
that are laid on the water singly, and (2) those that are laid so that
the eggs are adherent to each other and float in the form of a raft.
In the case of the single eggs these float with their length parallel
to the water-surface, and in the case of the raft-eggs they float in a
vertical position.
In both classes the individual eggs often have a higher specific
gravity than water, and consequently not being particularly buoyant,
if submerged and the adherent air detached, they sink to the
bottom. The membrane, owing to its water-proof properties, acts
as an attachment to the water surface film, causing the egg to float
as a dry sewing needle may float when it is placed so gently upon
the water that the surface tension of the film is not overcome by the
weight of the needle. Naturally, the substance of the egg weighs
far less than a proportionate bulk of steel, and the stability of the
* According to Nicholson the membrane has a highly specialised attachment to the chorion.
4*9
egg at the surface is thus far greater even in those eggs unprovided
with floats.
Nature has apparently recognised that in certain species a greater
stability is needed than that afforded to single floating eggs and
has developed the plan either of massing the eggs together to form
a boat-shaped raft, or by ballooning the enveloping membrane in the
single eggs to form the so-called ‘ floats 1 so commonly found among
the ova of the Anophelinae. The ‘ floats/ however, do not act to
any great extent as true floats giving buoyancy .to the eggs, but
support them on the surface film securely by presenting a larger area
on which the weight of the eggs may be distributed against the
surface tension. This .fact may be demonstrated by forcibly
submerging the eggs, when it will be found that they will either sink
or very slowly rise to the surface, generally failing to regain their
original position. Mosquito eggs that are laid in raft-formation are
almost impossible to submerge without breaking up the raft form
and detaching the eggs separately, on account of the air cells formed
between the adjacent sides of the eggs. All mosquito eggs are
submerged with difficulty, but submersion can be accomplished with
the aid of a fine earners hair brush by simply pressing on the eggs
with the brush until each egg is driven below the water surface.
THE OVA OF ANOPHBLBS MACUUPBNNIS
The average dimensions of the ova of A. maculipennis are,
length 075 mm., breadth across the widest part of the float
0*27 mm. Eggs from different batches vary in their dimensions
slightly, but the average variation is remarkably small. When the
eggs are viewed from above while they are floating normally on the
water it will be seen that the longitudinal outline of the dorsal surface
is roughly cigar-shaped and the floats which lie on each side give
the egg a slightly waisted appearance. The dorsal surface is
comparatively flattened, while the lower or ventral surface of the
egg is evenly rounded. One end of the egg is distinctly blunter
and broader than the other. On the ventral side, just below the
extremity of the broader end, the micropyle is situated, and it is
from the broader end that the larva emerges. The floats are placed
420
on either side of the egg almost at the middle point, though they
are actually a little nearer the more pointed end and extend for
rather more than a third of the egg's total length.
The float membrane is finely striated transversely, but the
striations do not indicate septa dividing the floats into compart¬
ments. The membrane which forms the float is superimposed upon
the main enveloping membrane, being attached to the main envelope
only along the ventro-lateral edges. This separate sheet of
membrane which forms the float is ballooned outwards and curved
upwards until it touches the dorso-lateral surface of the egg, to
which, however, it is not adherent.
By submerging an egg it can be viewed from several aspects.
Laid on its side it presents the outline shown in Plate XXII,
fig- 1 (A)> an< 3 upside down it has the outline shown in fig. i (r).
It will be noticed that the under surface of the egg is somewhat
boat-shaped and draughts progressively less towards the extremities
of the egg; also that the enveloping membrane on the under surface
of the egg shows hexagonal markings. The enveloping membrane
embraces the egg completely except at the micropyle, where a small,
round, black area is visible on the under surface of the blunter end
of the egg. When the upper surface of the egg of A . maculipennis
is examined it will be seen that the enveloping membrane spreads
out in areas of unequal thickness, presenting, in consequence, dark
and light markings as shown in fig. I (a). In these markings I find
a very easy method of distinguishing the eggs of A. maculipennis
from those of A . bifurcatus (compare figs, i (a) and 2 («))•
The markings are easily seen with a hand lens x 6, and to the
unaided eye give the eggs of A. maculipennis an ash-grey hue,
whereas the eggs of A . bifur catus, having no mottled markings on
the dorsal surface, appear almost black in colour. The advantage
of being able to distinguish the eggs of these two species from each
other is considerable, especially when the species are being bred in
the laboratory. Only one cage need be used, in which both species
may be kept together; and oviposition may be allowed to take place
• in the same dish. The ova are then easily separated.
421
THE OVA OF ANOPHELES BIFURCATES
Average length o*6i mm.; breadth across the widest part of the
float 019 mm. In general outline and structure the ova of
A. bifur cat us closely resemble those of A. ntaculipennis , but the
enveloping membrane which cradles the egg to the water line does
not extend over the dorsal surface,' and the dense black egg-shell is
naked. The floats in this species overlap the lateral edges of the
egg and give it a distinctly waisted appearance between the floats;
moreover, the floats are often of unequal size, and asymmetrical.
The two ends of the egg taper to a nearly equal degree, and the
curvature of the ventral surface is greater than in the case of the
eggs of A. ntaculipennis , causing the ends of the eggs to be more
conspicuously upturned.
THE OVA OF ANOPHELES PLUMBEUS
The average measurements are, length 0*56 mm.; greatest
breadth 017 mm. It has been found that to get A. plumbeus to
tolerate laboratory conditions is far more difficult than with
A. ntaculipennis and A. bifurcatus. It was only after numerous
attempts that I finally succeeded in getting 1 wild * specimens to
oviposit in the laboratory. These specimens were captured, in
Ejpping Forest by exposing ourselves to attack and allowing the
mosquitoes to feed. When replete, the mosquitoes were secured by
placing a glass tube over them as they rested upon the skin of the
arm. They were then transferred to the laboratory and placed
in large insect cages containing an almost moisture-saturated
atmosphere at a temperature of 30° C. Porcelain photographic
developing dishes containing the water from beech tree-holes on
which a few dead beech leaves floated were placed in the cages.
Four or five days afterwards the ova of this species were found
floating on the water contained in one of the dishes. Owing to the
colour of the tree-hole water and the form of the eggs they were
exceedingly difficult to see, and when viewed as they floated on the
deep brown-coloured water the ‘ floats * were quite invisible. The
eggs were carefully transferred to clean tap-water, and were then
seen to be particularly beautiful objects (Plate XXII, fig. 3).
Unlike the ova of A. ntaculipennis and A. bifur catus, the ova of
422
A . plumbeus are somewhat like the ‘ diamonds 9 of playing cards in
outline, and are completely surrounded by the floats. The eggs are
more pointed at the ends, both ends being alike, and bend
slightly upwards. The central part of the upper surface is also
raised, and the lower surface is evenly rounded.
It will readily be seen from the illustrations accompanying
this paper that the ova of A. maculipennis , A. bifurcatus , and
A . plumbeus are so conspicuously different from each other that the
differences afford an effective means of identifying the species.
FACTS CONNECTED WITH OVIPOSITION
Anopheles maculipennis .
There is no difficulty in getting the females of this species to
oviposit in the laboratory. Specimens captured in pigsties and
placed in a large insect cage in the laboratory live well, and will lay
several batches of eggs if a guinea-pig is placed in the cage overnight
two days after the first and subsequent batches of eggs are laid.
The eggs are always laid during the hours erf darkness; most often,
I think, in the early morning hours before sunrise. I have once
obtained a batch of ova at 12 o’clock noon by placing the cage in
a dark room.
The eggs when first laid are white, but they usually darken
rapidly. Sometimes individual eggs in a batch will, however,
remain white and fail to hatch, due, I think, to non-fertilisation.
This condition is most readily seen in the ova of females that have
been kept in the laboratory for some time and have laid numerous
batches of eggs, and it is probably to be explained by a diminished
supply of spermatozoa in the spermathecae.
The average batch of eggs laid by A . maculipennis in the
laboratory has been found to be about one hundred and fifty.
Anopheles bifur catus.
The eggs of this species are more difficult to obtain, for two
reasons. Firstly, the adults are only numerous in many localities
between the months of March and the end of May; and secondly,
while I have found it easy to get oviposition to take place in the
laboratory in the early months of the year, yet specimens captured
4*3
with difficulty after the end of May and placed under similar
conditions cannot be induced to lay, even after they have fed and
the ovarian eggs are fully developed. Differences in atmospheric
temperature seem to account for this, as I find that as soon as the
weather becomes cooler in the autumn and the mosquitoes become
more numerous again, the females will readily oviposit. The
average batch of eggs from A. bifurcatus in the laboratory has been
found to be smaller than that of A. maculipennis , comprising about
one hundred and twenty.
Anopheles plumbeus .
This species has been the most difficult from which to obtain ova.
As mentioned previously, repeated attempts had to be made with
fed specimens captured in Epping Forest before I was finally
successful. So far, I have only managed to obtain three batches at
different times from two females. In each case the batch was quite
small, consisting of from fifteen to twenty-one ova.
With one of the females the second batch was laid a day later
than the first, and this is probably a usual practice with the
species to avoid overstocking any one breeding-place, since the
majority of tree-holes have only a small water-holding capacity, and
comparatively few larvae are to be found in each.
I have recently extended my observations to the ova of the
British culicine mosquitoes, and find that in the ova of the species
that I have so far examined there are remarkable specific differences
sufficiently marked to be seen with the aid of a hand lens x 6, and
to enable the identification of the species to be easily made. On this
subject a further publication will be made shortly.
Probably owing to the difficulty in getting the majority of the
species of mosquitoes to oviposit under laboratory conditions, there
is a distinct scarcity of information on the specific differences in the
ova. Most of the work on the biology of a species in the existing
literature begins with descriptions of the larvae in the first instar,
and ends with descriptions of the adult and its life history. The
egg stage is, nevertheless, quite as important as any of the life
stages—in many ways, from a practical point of view, a particularly
important stage. Nuttall and Shipley (1901 and 1903), James and
Liston (1911), and Howard, Dyar and Knab (1919) are noteworthy
4*4
in having given special attention to the description of certain
Anopheline ova, but considering how numerous are the species of
the Cultcidae , it may be said that study and description of their ova
has been almost neglected.
REFERENCES
Giles, Geo. M. (1902). ' A Handbook of the Gnats or Mosquito/ J. Bale Danielsson Sc
Sons, London.
Howaxd, Dyar and Knab (1919)- * The Mosquitoes of North America and the West Indies/
Smithsonian Institute, Washington, D.C., U.S.A. Vol. I, p. 140, Vol. II, Plates
H5-7*
James, S. P., and Liston, Glen W. (19sx)« ‘A Monograph of the Anopheline Mosquitoes
of India,' p. 33.
Mitchell, E. C. (1907). ‘Mosquito Life.' Putnam Sc Sons, New York and London.
Nicholson, A. J. (1921). * On the Development of the Ovary and Ovarian egg of a Mosquito
Anopheles maculipennis , Meig. Quarterly Journal of Microscopical Science. VoL 65,
P. Ill, pp. 395-448-
Nuttall and Shipley (1901 and 1903). * Structure and Biology of Anopheles/ Journal
of Hygiene , pp. 49-81 and Plate II.
426
EXPLANATION OF PLATE XXII
Fig. 1. Ova of Anopheles maculipennis: ( a ) dorsal, ib) lateral,
(c) ventral aspects of ova.
Fig. 2. Ova of Anopheles bifurcatus: (a) dorsal, (b) lateral,
(c) ventral aspects of ova.
Fig. 3. Ova of Anopheles plumbeus: ( a ) dorsal, (b) lateral,
(c) ventral aspects of ova.
PLATE XXII
&? Paraiitol., I’ol. XI
4*7
THE TRYPANOCIDAL EFFECT OF
PHENYLGLYCINE AMIDO ARSENATE
OF SODIUM ON T. BRUCEI IN RATS
AND T. RHODESIENSE IN MICE
BY
S. ADLER, M.B.
(Received for publication 23 November, 1921) .
This drug* is a white amorphous powder readily soluble in
distilled water, and yielding in 5 and 10 per cent, concentration
a perfectly clear, colourless solution. On standing, however, for
some days, a yellow colour develops, whether the solution is kept
exposed to light or in the dark. A 5 per cent, solution when
exposed to light became yellow in seven days; when kept in the
dark, in six days. Daily sterilizing for ten minutes did not prevent
the development of the yellow colour, but rather accelerated it.
Rats . Minimum lethal dose . In Table I are set out the experi¬
ments performed in order to ascertain the minimum lethal dose for
rats; in all cases the drug was used in a freshly prepared solution of
5 or 10 per cent., and was injected intraperitoneally; it produced
no irritation. Healthy and infected animals were used in this
experiment.
It will be seen from the table that no dose was toxic to the
animals injected until the amount of V 2 gms. per kilo was attained.
Relationship of Toxicity to age and change of colour of solutions .
The number of experiments bearing on this point is rather
limited, but serves to show that the toxicity increases on standing,
e.g ., although the minimum lethal dose of freshly prepared solution
for rats proved to be i*2 gms. per kilo body weight, an animal
which was injected with 0*18 gms. per kilo of a twenty-four hour
old solution followed on the next day by a dose of 0 36 gm. per
kilo of a forty-eight hour old solution, died after severe symptoms
of poisoning on the sixth day after the second injection. In the
Kindly put at our disposal by Messrs. May & Baker, LtiL, Battersea, London.
428
Table I.
Showing the minimal lethal dose for rats.
No. of
Experiment
Dose in gms., per kilo
Infected or healthy
Remarks
i
o *33
. Healthy
No toxic effects
2
0*46
Healthy
No toxic effects
3
°’43
Healthy
| No toxic effects
4
o'67
Infected
i
No toxic effects
5
o-68
Infected
1 No toxic effects
i
6
07
Infected
!
| ‘ No toxic effects
7
071
Healthy
1
No toxic effects
8
075
Infected
No toxic effects
9
0*82
Infected
No toxic effects
IO
o’yo
Infected
No toxic effects
11
ro
Healthy
1
No toxic effects
12
ri
Healthy
No toxic effects
• *3
1*2
Infected
Animal died in 22 hours
14
r 4
Healthy
Died on 4th day
4*9
above solution, although highly toxic, no change in colour was
evident. Further experiments proved that in still older solutions
the toxicity diminished, e.g., on the eleventh day, when the
solution was deeply yellow, three animals were each given a dose,
approximately equal to the combined doses given above, without
producing any symptoms. The first of these three animals was
injected with a solution which had been kept in the daylight and
boiled each day for ten minutes, the loss from evaporation being
made up by addition of distilled water to the original volume before
injection. The second was injected with a solution which was
unboiled and kept in daylight; the third with solution unboiled and
kept in the dark.
The increase in toxicity which was evident in the forty-eight hour
old solution, was accompanied by a definite increase in trypanocidal
power, for after injection of 0 36 gm. per kilo, .trypanosomes
disappeared from the peripheral blood within twenty hours, and
were absent from the blood until the time of death. Examination
for trypanosomes of the organs by smears proved negative, as did
also the injection of emulsion of organs and blood into healthy rats.
A similar sterilizing effect could only be produced by much larger
doses of the freshly prepared drug, e.g ., a dose of 0 67 gms. per kilo
in freshly prepared solution injected into a rat at approximately
the same stage of infection as the previous one, only caused the
disappearance of the parasites from the peripheral blood in forty-
eight hours, and did not prevent their re-appearance twenty-four
hours later.
Toxic effects of freshly prepared solutions.
No toxic effect was observed below the dose of Y2 gms. per
kilo; a heavily infected animal died within twenty-four hours of
receiving this dose. The signs of poisoning noted before death
were blindness and refusal to take food. No gross haemorrhages
were found after death, but the whole intestinal tract showed
numerous minute haemorrhages. No trypanosomes were found in
the peripheral blood, either immediately before or after death.
Toxic effects of old solutions.
Toxic effects were noted after a dose of 0 36 gm. per kilo,
but not until four days had elapsed. On the fifth day the
430
animal appeared ill, lying curled up, breathing irregularly and
spasmodically, it refused food, was blind, and staggered in its gait
when moved. Haemorrhages were observed from the conjunctiva,
anus and urethra. Post-mortem, the whole intestinal tract was
found to be haemorrhagic, the liver was enlarged and soft, and
showed yellow mottling; the kidneys were very soft, but not
enlarged nor dark in colour; the spleen showed yellowish patches.
Minimum . curative dose .
In Table II are set out the experiments performed in order to
demonstrate the minimum curative dose in rats.
Table II.
Shewing effect on rats infected with T. bructi.
Experi¬
ment
Trypanosomes
per field.
Ocular 4,
Obj.J
Dose in gms.,
per kilo
Day of
disappearance of
trypanosomes
from peripheral
blood
Day of
re-appearance
Remarks
i
4 °
°*33
1
5
2
Swarming
0*67
2
3
...
3
Swarming
o*68
1
2
4
Swarming
°*7
1
...
No relapse
after
7 months
5
36
075
1
...
No relapse
after
7 months
6
0*82
1
...
No relapse
after
7 months
7
25
0*91
1
...
No relapse
after
7 months
8
Swarming
1*2
Died in 22
hours. No.
trypano¬
somes
Control
Swarming
0*17
(atoxyl)
1
>7
It will be seen from the above table that while the atoxyl control
animal was rendered free from trypanosomes in its peripheral blood
+ 3 *
Tabl« III.
Showing effect on mice infected with T. rbodenense .
Experi¬
ment
Trypanosomes
per field.
Ocular 4,
Obj. }
Dose in gms^
per ldlo
Day of
disappearance of
trypanosomes
from peripheral
blood
Day of
re-appearance
Remarks
i
Swarming
o *5
2
4
...
2
o*6
2
6
...
3
Swarming
074
2
5
...
4
Numerous
0*83
1
8
...
5
0-9
2
5
6
Swarming
o # 9
*
4
7
Numerous
ro
5
...
8
Swarming
VI
2
...
Alive and well
5 months
iater
9
Numerous
1*25
2
4
...
IO
Numerous
r6
1
6
...
ii
Numerous
2*0
1
12
...
12
Swarming
**5 ,
2
7
*3
Control
i
3 *o 1
i
. 1
...
Died after
3 d»y»
43 2
by 017 gms. atoxyl per kilo, but relapsed in seventeen days, the
experimental animals were rendered free from trypanosomes when a
dose of 0 7 gms. per kilo of phenylglycine amido arsenate of sodium
was reached, and did not relapse.
Action in Vitro.
No trypanocidal action in vitro was observed, either by the drug
itself 01 by the serum of animals, twenty-four hours after they had
been rendered free from trypanosomes by injection of the drug.
Mice. Minimum Lethal Dose.
In Table III are set out the experiments performed in order to
demonstrate the minimum lethal dose for mice and the effect on
T. rhodesiense .
It will be seen from the table that the minimum lethal dose was
3 gms. per kilo; the effect on the trypanosomes was negligible up
to a dose of 2*5 gms. per kilo, only one animal failing to relapse.
SUMMARY
Phenylglycine amido arsenate of sodium can be used in freshly
prepared solutions in distilled water for intraperitoneal injection
into rats and mice; the solutions on standing become toxic, and later
become yellow in colour.
For rats, the minimum lethal dose of the freshly prepared drug
proved to be V 2 gm. per kilo of body weight.
For rats infected with T. brucei , the minimum curative dose is
0 7 gm. per kilo of body weight.
In vitro the drug has no appreciable action on trypanosomes, nor
has the blood of treated animals immediately (twenty-four hours)
after becoming trypanosome-free.
The drug has no curative effect on mice infected with
T. rhodesiense.
A remarkable feature of this drug is its relatively high minimum
lethal dose. Although the drug contains 26 per cent, arsenic, the
minimum lethal dose was found to be 12 gms. per kilo for rats and
3 gms. for mice.
I am indebted to Professor Blacklock for carrying out the
necessary inoculations for the experiments.
433
NOTE ON BISMUTH AS A TRYPANOCIDE
BY
S. ADLER, M.B.
(Received for publication 23 November , 1921)
The effect of bismuth in the form of soluble bismuth sodium
tartrate in solutions of various strengths was tried on animals infected
with T . rhodesiensc and T. brucei {Nagana ferox) respectively.
Using this drug, the minimum lethal dose for healthy mice was
found to be 0*047 grammes of bismuth, and for healthy guinea-pigs
0*062 grammes of bismuth per kilo body weight.
In animals which died after injection of bismuth sodium tartrate
deposits of bismuth were found in all cases in the liver, frequently
in the spleen, and less frequently in the kidneys.
T. rhodesiense .
The minimal lethal dose cleared the blood of trypanosomes in a
mouse within twenty-four hours, but the animal died in two days.
Any dose below this failed to clear the blood of trypanosomes.
T . brucei {Nagana ferox).
Although the drug cleared the blood of trypanosomes in guinea-
pigs, yet relapses occurred in a few days, thus : —
Dose per kilo, body weight
Injected
Trypanosomes re-appeared
o*o 18
1
j 20.4.21
1 28.4.21
0*022
j 26.4.21
2.5.21
0*024
1
9.5.21
18.5.21
0*032
9.5.2 1
1
21.5.21
In no case was a cure obtained.
I am indebted to Professor Blacklock for carrying out the
necessary inoculations for these experiments.
435
MALARIA ON A VENEZUELAN
OILFIELD
BY
J. W. W. STEPHENS, M.D., F.R.S.
(Received for publication 5 December, 1921)
PLATES XXIII AND XXIV
The oilfield is situated on the eastern side of Lake Maracaibo,
Venezuela, about 80 miles from the head of the lake, and 10 miles
inland from its shores. The following observations were made in
the month of August, 1921.
MALARIA IN THE NATIVE POPULATION
The extent to which malaria prevailed in the native population
of various villages in the district is shown in the following table: —
Age Period.
0—10
i
11
1
—20
21
—
No.
exam.
Percentage
infected
No.
exam.
Percentage
infected
No.
exam.
Percentage
infected
Old Village .
10
10
8
ir 5
0
1
—
New Village .
-
32
468
39
23*0
49
8 -i
San Pedro.
9
66*6
6
0*0
1
0*0
Los Barrosos .
9
0*0
9
0*0
0
—
San Timoteo .
2 7
37
4
0*0
0
—
The relationship of malaria in the Old Village and New
Village respectively to malaria among the white employees on
the oilfield we shall discuss later. The other three villages were
not in the immediate neighbourhood of the oilfield, but it may be
pointed out here that the absence of infection in Los Barrosos was
probably due to the fact that anophelines were only found at a
distance of half a mile, and then only in one small clay-pit about
436
12 feet square. San Timoteo is a village built on piles over Lake
Maracaibo. Its low endemic index, 37 in children under ten, is
thus probably due to its position over the lake and to the fact that
only one small anopheline larva was found after prolonged search on
the mainland. San Pedro is four or five miles from the oilfield, and
its high rate is probably due to its being situated close to extensive
anopheline swamps.
SPECIES OF PARASITES FOUND
In over two hundred blood specimens examined, malignant
tertian malaria (crescents') was only found twice, simple tertian and
quartan parasites were about equally common, a peculiarity of
infection which I had not previously seen in Africa or India. The
frequency with which pigmented leucocytes were observed in the
‘ positive * films was a noteworthy feature.
MALARIA AMONG THE WHITE STAFF
No precise information was forthcoming as to the nature of the
sickness among the white employees (about thirty in number). An
examination of the blood of sixteen employees in apparent good
health revealed nothing. In the case of one employee suffering
from fever, simple tertian parasites were found. In the absence of
definite records, one had therefore to assume that the sickness
prevalent among the staff was due to malaria.
POSITION OF THE STAFF HOUSES
They are situated (vide Map), (1) on two ridges a few hundred ,
yards from the first source of Anopheles , viz., the small anopheles
swamp in the Asphalte creek, the houses on the easterly ridge actually
overlooking the swamp. (2) From the second source of Anopheles
the ‘ Anopheles Swamp * which begins due west and even north-west
of the New Village, they are situated less than half a mile. (3) They
are situated less than a quarter of a mile from a source of malaria
infection, the Old Village, and (4) since the year 1920 half a mile
from a second source of infection, the New Village.
We have here then an example of the conditions so frequently
Annals Trop. Med . & Parasitol, , Vol. XV
PLATE XX111
C. Tinling A* Co., Ltd., Imp
Hoof rnarU
437
founds in the tropics, namely, native villages and a source of
Anopheles in close proximity to a white population, and here, as
elsewhere, the conditions would completely explain the existence
of malaria among the white employees.
THE ANOPHELINES
The district under investigation can be divided into four
areas: —
A. The Jungle.
B. The cleared area.
C. The residential and Old Village area.
D. The New Village area.
The relative positions of these will be seen on the map.
Area A. The Jungle.
Numerous examinations were made of all discoverable water in
the jungle so far as it was penetrable. In many cases, pools and
streams and * tatucales/* apparently favourable breeding-places,
were examined with negative result, often, no doubt, due to the
presence of abundance of small fish (in some cases due to a covering
of Lemma (duckweed). In other cases, however, jungle pools were
1 Culex 9 positive.
One interesting example was encountered, viz., that of a borrow-
pit, where the snouts of dozens of small fish could be seen voraciously
hunting the surface for food, and yet ‘ Culex 1 larvae in abundance
were found in the protecting weed and rubbish at the sides, but
none away from there.
Again where the jungle was apparently dry over large areas,
yet 1 Culex * was present and bit freely in the daytime when one stood
still.
Anopheles (mosquitoes or larva) was, then, absent from the
jungle at least in August, and mosquitoes caught in the jungle at
long distances from the oilfield were invariably ‘ Culex *
Area B. The Cleared Area.
Here and there a few * Culex 9 breeding-grounds were found,
generally in small numbers. A swamp a hundred yards or so long,
• Tatucales: Diminutive islands or mounds intersected by water, and in which progress could
only be made by a series of jumps.
+ 3 *
passing in front of the native mechanics* quarters i to 6 and close to
the next area, is of interest in that repeated search showed it to be
Anopheles negative, but it contained a small 4 Culex 9 area near its
origin. It had been already pafrtly, but incompletely drained, and
fish abounded.
Area C. Staff Houses Area.
The Staff Houses arc situated for -the most part on two ridges
themselves separated by a dry ravine. These ridges are bounded
on the N.W. and N.E. by the Mene Grande and Asphalte creeks
respectively. These creeks merge at their origin northwards, and
are bounded by the slopes of the ridge on which the Star Water
tank is situated. Again further north to the west and the east
this ridge descends in sloping ground, dry for the most part and
covered with asphalte until in a N.W. direction the creek ‘ Piedritas
blancas 9 is reached.
The whole of this large area is for the most part dry. What
little water there is, is found in the slopes draining to the road from
the Old Village to Zo, but frequent examinations of all collections
of water in this area were negative. As regards the Asphalte creek
itself, this was also negative except for a small swamp some fifty
yards long in its upper portion. Here Anopheles larvae in scanty
numbers were found. The rest of the creek had already been
drained; the ditches contained numerous fish and considerable
quantities of oil. The swamp above referred to was caused by a
small depression in the surface, and was easily abolished by cutting
the.long grass and filling with earth from the adjacent higher
ground.
Area D.—or New Village Area.
The New Village is situated on the Sabana de Matajey de Raya
on elevated ground (thirty metres) a hundred yards or more from the
creek El Mene or water creek. From the elevated flat ground on
which the village lies a number of dry ravines lead down to a swamp
some fifty yards or so in width, bordering the stream formed by the
junction of the streams from the Asphalte and El Mene creeks. This
swamp extends along the edge of the Sabana in a curved course
(probably as far as San Pedro), and was found to contairr Anopheline
larvae—always, it is true, scanty in number—for a distance of,
439
roughly, about three-quarters of a mile. Although fish were
plentiful in the two drains that had already been cut in part of it,
but which were blocked when examined by me, yet Anopheles were
present always in small numbers, but from its extent this swamp and
its tributary swamps ( vide infra) formed the main source of
Anophelines . The Tatucales, out in the Sabana, contained fish,
and was negative.
The ravines below the village to which reference has been made,
extend in a south-easterly direction, becoming less steep and with
flatter bottoms so that they are no longer dry but are occupied
by swamps; it is these secondary tributary swamps and the
main swamp which they eventually join, which formed the great
Anopheline breeding-ground.
In the four areas examined, Anopheles were found breeding in
two only, and in one of these only in a swamp of small dimensions.
The breeding-grounds were thus in August definitely restricted;
Anopheles were not breeding in the jungle nor in the open cleared
area, nor in the waters that existed on the asphalte slopes of
Area C.
The swamp in which Anopheles were found breeding was over¬
grown with a variety of grasses, some up to 6 ft. high; in other parts
it more resembled marshy ground with short rushes in part trodden
by cattle; in other parts again ‘tatucales* formation. In no case
were Anopheles found amidst overhanging trees.
Anopheles (C. argyrotarsis) were also found breeding in one of
half a dozen clay borrow-pits some three miles away from the camp.
Adult Anophelines.
The search for anophelines in the native huts in.the daytime
was completely fruitless, and culicines also were very scanty. This
condition was in marked contrast to those observed by me in the
neighbourhood of Lake Valencia, which I visited on my way home,
where in the daytime, in the verandah of a hut, it was easy to collect
numerous anophelines, embracing three different species. On the
oilfield itself I was only able to secure by capture at night in the
native village sixteen specimens of C. argyrotarsis , and during my
stay in the camp no anophelines and very few culicines were seen by
440
me in the house I occupied. It should be stated that the windows
of all the houses in the camp were wire-screened, and mosquito-nets
were in general use.
PROPHYLAXIS
The existence of infection in the Old Village and the absence of
Anopheles from any area nearer than an isolated small, grass-grown
swamp in the Asphalte creek and the great Anopheline Swamp
adjacent to the New Village proves, I think, that we have a normal
flight range of about half a mile.
We may consider prophylactic measures under the following
headings: —
(a) DRAINAGE OF ANOPHELINE SWAMP.
Towards the end of my visit, 100 yards or so had been cleared
of grass and existing choked drains cleared and additional ones,
constructed, with the result that the swamp rapidly became dry and
free from larvae in this section. I suggested that the swamp should
be drained for about a mile. The efficacy of this measure can be
estimated by observing the effect (i) on the number of anophelines
caught in the New Village month by month; (2) on the endemic
index of the New Village; (3) on the sickness rate of the white staff.
If the drainage results in a complete or almost complete suppression
of anophelines, further measures would hardly be necessary, but in
the event of this not being so, other measures would have to be
adopted.
( b ) Removal of the Old Village.
This source of infection in close proximity to the white staff
quarters should be abolished, and if further protection is sought
various stray native huts which exist here and there should also be
removed.
( c ) The New Village.
If drainage of the anopheline swamp is unsuccessful, it would
be necessary to consider the question of removal of the New Village
to an area free from anopheline breeding-places; such a one existing
a mile or so from its present position. For elimination of malaria
from the village itself, quinine administration is advisable; and
further, if it remains in its present position wire screening of the
huts could be advantageously employed, without prohibitive cost.
(d) A more fundamental and expensive - procedure would be the
removal of the white staff houses. An ideal site for these exists on
the Star Tank ridge, at an elevation of about 300 ft. and at a
distance of about a mile from the New Village and Anopheline
Swamp.
One sees, accordingly, that the existing conditions have arisen
from a lack of appreciation of the fundamental principles responsible
for the infection of the white man amidst a native population,
namely, dose propinquity to that population, with a supply of
anophdines also adjacent. Had one been laying out the camp
de novo, the following arrangement would, I believe, have led to the
protection not only of the white but also of the native population: —
*(1) The white staff houses should have been placed on the Star
Tank ridge; (2) the native village should have been placed a mile
from this site and also a mile from the source of anophelines.
The conditions prevailing were particularly favourable for such
an arrangement, namely, (1) the complete absence of anopheline
breeding-grounds over a large part of the area considered, and
(2) the large tract of ground in which collections of surface water of
any sort were absent (in August).
NOTES
The oilfield was situated in Lat. 9 0 .75' N. and Long. 71 0 .15' W.
The indoor temperatures in August ranged from 85°-8(y5 0 F. maxima
to 75°-8r5°F. minima. Heavy rain lasting about half an hour fell
towards evening on five out of twenty days.
The only species of Anopheline found in the area was Cellia
argyrotarsis, distinguished by having three and three-quarters hind
tarsi almost completely white. The palmate hairs of the larva of
this species are variable. Usually there are palmate hairs on
segments two to seven, that on segment two being small and hard
44 2
to see. Examples were also found with well developed palmate
hairs on segments one to seven, together with a well developed
thoracic palmate hair.
Examinations of sixteen specimens of C . argyrotarsis for sporo¬
zoites proved negative, but Tovar states that this species transmits
in Venezuela.
The small fish which abounded in the small streams and in the
swamps of the district belonged to the following species: —
Gambusia ( Poecilid ) tridentigera ,
Haplochilus sp., and
Chromides (.Acara ) dorsigera.
EXPLANATION OF PLATE XXIV
Fig. i. Old type of Bungalow.
Fig. 2. Modern type of Bungalow.
Fig. 3 * The Peons’ Cock-pit.
Fig. 4. Bird’s-eye view of Staff houses. The ridge with Star
Water Tank in the distance.
f ig* 5 - The Asphalte slopes. The arrows point to two asphalte
mounds.
• •Ill 1
* ' • i
j : . LI 1 1
1 |
T K Y ^
U _ t 1
_1_L
^ a/
S' :
[<f«K» m '0
tmmziy
KJk-vi-S
lj ^ • <
■ 1
445
NOTES ON CULICIDAE COLLECTED IN
VENEZUELA
BY
A. M. EVANS, M.Sc.
{Received, for publication 25 November, 1921)
Plates XXV and XXVI
During the course of the investigations in Venezuela, recorded in
the foregoing paper, a number of mosquitoes were collected by
Professor Stephens at Mene Grande, which is situated about ten
miles inland from Lake Maracaibo. Specimens were also caught at
Maracay near Lake Valencia, which is separated from Lake
Maracaibo by a range of mountains, and on the Island of Curasao.
The following is a list of the species collected at each of the three
places:—
Lake Maracaibo
Anopheles argyrotarsis, R.D.
Hatched from larvae taken in swamp A <J 3, 9 9 6. Mene Grande,
22.8.21, $9 5.
Aedes serratus, Theo.
Mene Grande, 3.8.21, 9$ £. Biting in jungle, Mene Grande,
• 5.8.21, 99 6. Caught in jungle, biting horse and man,
5.8.21, 9 1; 11.8.21, 99 11.
Aedes scapular is, Rond.
Mene Grande, jungle, 11.8.21, 99 3-
Taeniorhynchus titittans. Walk.
Mene Grande, biting in jungle, 99 3-
Culex (Neomelanoconion) ckrysothorax, Newstead and Thomas.
Reared from pupa taken in swamp, Mene Grande, 7.8.21, 9 1.
44 6
*Culex ( Culex) coronator, D. and K.
Mene Grande, 6.8.2 r, 33 4, 9? 6. Hatched from larvae, Mene
Grande, 1.8.21, 33 2 ; 11.8.21, 99 2; 33 3, 9 1.
*Cidex (Culex) nigripalpus, Theo.
Mene Grande, 3 1.
Psorophora posticata , Wied.
Mene Grande, 11.8.21, 9 ? 2.
Joblotia digitatus, Rond.
Jungle, Mene Grande (about 20 miles inland from Lake
Maracaibo), 20.8.21, 99 2.
Maracay
(about 10 miles from Lake Valencia)
Anopheles argyrotarsis, R.D.
Maracay outskirts, 8.9.21, 9 1.
Anopheles albimanus , Wied.
Maracay outskirts, 8.9.21, 3 1, 9 1.
Anopheles albimanus var. tarsimaculatus , Goeldi.
Maracay outskirts, 8.9.21, <? 1, 9 1. Maracay outskirts, 5
specimens.
Anopheles pseudopunctipennis, Theo.
Maracay outskirts, 7 specimens.
Aedes scapular is , Rond.
Maracay outskirts, 8.9.21, 9 1.
Aedes trivittatus, Coq.
8.9.21, 99 2.
*Culex (1 Culex) quinquefasciatus , Say.
Maracay outskirts, 8.9.21, 33 3-
* Culex (Culex) virguUus , Theo.
Maracay outskirts, 8.9.21, <? 1.
Psorophora posticata , Wied.
Maracay outskirts, 8.9.21, 9 1.
Determined by the male genitalia (See Dyar, 1918V
447
Curacao
Aedes (Stegomyia) fasciola, Fabr.
Breeding in tub, Ice Factory, Curayao, 1.9.21, 2 1
*Culex ( Culex ) quinquefasciatus , Say.
From larvae breeding in tub of ice manufacturer, 30.8.21.
<? $ 9, 2 2 11; 1-2.9.21, $ $ 14, 2 2 31.
THE MORPHOLOGICAL CHARACTERS OF
ANOPHELES ARQYROTARSIS , R.D., AND A. ALBIMANUS f WEID.
Larvae. In their Monograph, Howard, Dyar and Knab (1917),
state that palmate hairs occur only on abdominal segments two to
seven in A . argyrotarsis, but that in A. albimanus there is an
additional small pair on the first abdominal segment. The larvae
of A. argyrotarsis , which Professor Stephens collected from the
swamps around Mene Grande, however, all possessed these
structures on the first segment of the abdomen. In some specimens
these were in a reduced and incomplete condition, but in a certain
number they were well developed, and in these latter specimens a pair
of palmate hairs was also present on the thorax, a position in which
they do not appear to have been recorded hitherto in either of these
species. The distribution of these structures is thus shewn to be
variable in the t;wo species, and cannot be used as a specific character
for determination of the larvae.
Male hypopygium . In view of the close resemblance between
the adults of these species, it appeared desirable to compare the
detailed structure of the male hypopygium. The figures in the
Monograph of Howard, Dyar and Knab (1912) tend to exaggerate
the differences present, and it is obvious that the morphology
was not clearly understood. Christophers (1915) described the
hypopygium of A. albimanus , and placed in one group with this
species A. argyrotarsis , A. .tarsimaculata, and A . bellator. This
group was based on the number and arrangement of the large spines
arising from the side-piece, but Christophers stated that it was
imperfectly studied. Edwards (1920) referred to the structure of
the mesosome (theca) of A. argyrotarsis , saying that it approached
nearest to the simple form seen in Ochlerotatus.
•Determined by the male genitalia (iee Dyar, 1918).
44 *
Before comparing the hypopygium of the two species, it is
necessary briefly to discuss the structure of certain of the constituent
parts. The nomenclature employed is that proposed by Edwards
(1920). The mesosome (theca) of A. albimanus has been described
and figured as ‘ massive and clubbed.' This appears to have been due
to confusion of the mesosome with a wide median membraneous lobe
(fig. 2 A and B, tn.l .) which arises from the membrane at the base of
the side-pieces on their upper (by rotation) sides. The position of
Fio. 1. <$ Hypopygium of A . argyrotarsu from below; tenth sternite not shown.
cl.y claspette ; mesosome ; s.p., side-piece ; ix, stalked spine.
this lobe in relation to the side-pieces is shewn in the drawings
(fig. 2 A and B). Distally it becomes bilobed, and the two halves
are produced downwards (towards the tenth stemites) so as to
embrace the distal half of the mesosome. So intimately are the
distal portions of the lobe associated With the mesosome, that in
mounted preparations it is often impossible to distinguish the
separate structures. When examined floating in oil under a
binocular microscope, however, and arranged so that a terminal view
449
is obtained the mesosome can be seen lying ensheathed on three sides
by the median lobe, and its downward processes. The separate
parts can then be dissected away and isolated. The form of the
mesosome and median lobe afford what appear to be reliable
O l millimeters J
Fio. 2. Bate of $ Hypopygium from above; mesosome and tenth stemite not
shown; drawn from specimens macerated in K.O.H. A— A. argyrotarsis. B— A. albimanus
var. tarsimaadata. mi., median lobe ; s.p., side-piece.
characters for the separation of the species A. argyrotarsis and
A. albimanus . The latter species and its variety tarsimaculata
appeared not to differ to any marked degree in the characters of the
hypopygium.
450
The Mesosome (fig. 3 A and B). It was found that the mesosome
can be best studied in specimens stained with carbol fuchsin. In
both species the halves of the mesosome are elongate plates, each
articulating basally with the chitinous pieces (j>.b.p .), which
represent the parameres and basal plates of other Culicidae.
The halves ( h ) approach each other on the upper side, but
below are connected by thin membrane which distally becomes
chitinised, forming a rounded apical plate ( [a.p .) concave above, and
membraneous at its margin.
In A. argyrotarsis the halves of the mesosome may be actually
contiguous for part of their length, causing the mesosome to be
tubular in this region. At the distal extremity of each half arises
a flat recurved spine (/.), toothed on its outer side, which is obviously
equivalent to the leaflets of the more specialised Anophelini.
Although the leaflets of other species are usually directed more or
less distally, Swellengrebel (1921) figures some recurved ones in
A. ( Myzorhynchus ) barbirostris , v. d. Wulp var.* pallidus , Sw.,
and in A. ( Myzorhynchus ) umbrosus , Theo. The greatest breadth
of the apical plate is greater than its height above the apices of the
halves of the mesosome.
In A. albimanus and its variety tarsimaculata , the halves of the
mesosome (fig. 3 B) have only a slight tendency to form a tube.
Leaflets are absent, and the greatest breadth of the apical plate is
4 $i
equal to or less than its height above the apices of the halves of the
mesosome.
The Median lobe . In A. argyrotarsis (fig. 2 A), the distal
portions Qf this structure are composed of a number of parallel
divisions, the exact form of which is exceedingly difficult to
determine in macerated specimens, owing to the extreme thinness of
the membrane, and its liability to distortion. There are no
long hairs or setae present. In A. albimanus and its variety (fig. 2 B),
the surface of the median lobe is thrown into a large number
of shallow folds distally, these folds involving the lateral descending
portion as well as the upper surface. The sides are thickly clothed
with long hairs, and a number of very delicate hairs occur on the
surface of the basal portion of the median lobe.
Side-pieces . No well marked specific characters could be found
in the large spines borne on the ‘internal surface of the side-pieces.
The pedicels of the stalked spines (fig. I, s.s .) are slightly shorter in
A. argyrotarsis than in A. albimanus . The claspettes . are in the
former species broad and not well differentiated from the surface
of the side-piece. In this they differ from the separate finger¬
like condition in which they sometimes occur in A. albimanus
(Christophers, 1915). In some specimens of the latter species,
however, they occur in the same condition as in A. argyrotarsis.
The tenth sternite is almost identical in the two species. The
appearance varies according to the position of the .paired chitinous
arms.
The main differences, then, between the male hypopygium of
A. argyrotarsis and A. albimanus lie in the form of the mesosome
and the membraneous structure here referred to as the median lobe.
The character of the mesosome of A. argyrotarsis , with its single pair
of leaflets and tendency to form a complete tube, suggests a
transitional stage between the generalised condition seen in
A. albimanus and the completely tubular form with numerous leaflets
which odcur in most species of the genus.
REFERENCES
Christophers (1915). Ini. Jovm. Med. Res. Vol. Ill, p. 379.
Dyar (1918). Ins. Ins. Mens. VoL VI, p. 94. Plates III and IV.
Edwards (1920). Ann. Trop. Med. & Parisitol . VoL XIV, p. 23.
Howard, Dyar and Knab (1912), (1917). Mosquitoes of North and South America and
the West Indies. Vol. II, Figs, 258, 263, 264. Vol. IV, Table on p. 967.
Swkllkngrzbkl (1921). Overged. uJ>. Tijds. v. Ent n Deel LXIV, 1921, pp. 39, 40.
EXPLANATION OF PLATE XXV
Wing of Anopheles albimanus var. tarsimaculata.
End of hind tarsus of A . albimanus var. tarsimaculata.
End of hind tarsus of A. argyrotarsis.
0 5 millimeter
454
EXPLANATION OF PLATE XXVI
Fig. i. Wing of Anopheles punc/ipennis.
Fig. 2. Wing of A. pseudopunctipennis.
Annals Trop. Med. ParasitolJ'ol. XT
PLATE XXVI
Fic.
1 millimeter
Fig. 2
A. M. ExcnSy ad. tint. del.
C. T ini ins Ltd., http.
4SS
A NOTE ON THE SYNONYMY- OF THE
GENUS ZSCHOKKEELLA, Ransom, 1909,
AND OF THE SPECIES Z. GUINEENSIS ,
(Graham, 1908)
BY
T. SOUTHWELL
AND
P. A. MAPLESTONE
{Received for publication 29 November, 1921)
In Simpson’s Report on Plague in the Gold Coast in 1908, Graham briefly
described a parasite from Cricetomys gambianum, to which he gave the
name Davainea (guineensis, n.sp, ?).
We have obtained from the Gold Coast on several occasions, through
the kindness of Dr. J. W. S. Macfie, large collections of worms, both from
C. gambianum and Epimys ( Mus ) rattus. The examination of this material
has led us to make the following observations:—
Beddard (1911, a), apparently unaware of Graham’s paper, redescribed
the same worm under the name Thysanosoma gambianum. Beddard
(1911, b) changed this name to Thysanotaenia gambiana, and again (1912)
to Zschokkeella gambianum. Baylis (1915) described Z. muricola, n.sp.,
from Epimys {Mus) rattus, and Meggitt (1921) described a new species of
Inermicapsifer (/. Zanzibarensis), from C. gambianum.
We have been unable to find any constant or adequate points of
difference between our material and the above three species; this is
clearly indicated in the table below.
For these reasons we have come to the conclusion that the worm
Z. guineensis has the following synonymy, viz.: Davainea {guineensis,
n. sp. ?) Graham, 1908; Thysanosoma gambianum, Beddard, 1911; Thysano¬
taenia gambiana, Beddard, 1911; Zschokkeella gambianum, Beddard,
1912; Zschokkeella muricola, Baylis, 19x5 and Inermicapsifer Zanzibar ensis,
Meggitt, 1921.
We agree with Meggitt (1921) that the genus Thysanotaenia should
lapse, and that no valid distinction has yet been made between the genera
Zschokkeella and Inermicapsifer. We therefore suggest that the genus
Inermicapsifer, Janicki, 1910, be likewise suppressed, and all the species
in it included in the genus Zschokkeella, Ransom, 1909, on account of its
priority.
456
Z. gambianum , Beddard Z. muricola , Baylis
/. zanzibarensis , Meggitt
Z. guiumu*
Genital Pore
About the middle. Anterior.
Afore—Shown anterior
in figure.
In anterior quarter.
Varie* from the artrncf
to about the middk,-
the same itrobik
Testes ...
In two separate groups.
(This is only implied in
the description and
is not definitely
stated).
In two groups, but almost
continuous.
Continuous; but fewer
in centre of segments.
May be in two
groups or these mar *
united by a hridst i
varying numben ti
testes, in the tm
strobila.
Vesictda seminalis
? Present. If so, repre¬
sented by a pouch near
the ovary.
Not seen.
Small and inside cirrus
pouch.
Small and inside arras
pouch. No extend
vesicula seminalis m
Ovary ,
May be regarded as
paired; partly separ¬
ated by yolk gland.
Single, fairly compact,
lobulated, crescentic.
Single, made up of strag¬
gling lobes, long,
slender and only slightly
connected.
A pear-shaped-erpa as¬
sisting of dub-ii?p;c
acini, with ritelhr
glands lying ck«W
posterior.
Receptaculum seminis
Long and
swollen.
not much J Small rounded organ near
ovary.
Large and thin walled.
Note —This structure
appears to be what
is usually considered
as the vagina.
As in Z. nuriala
No. of eggs in capsules
A few.
About 20.
One, but 9 to 10 eggs in
clusters.
From 3 to about 20.
Not *—Apparently a
egg capsdesfint «■
tain about ao
and in liter fab-
ment these ab|
divide into cspdei
containing kv
# Diagnosis compiled from observations on our materia] and from specimens which Dr. Meggitt
was kind enough to send us.
REFERENCES
Baylis, H. A.- (1915). A new Cestode of the Genus ZscbokkeeUa. Ann. & Mag. Nat. Hist*, Ser. 8,
Vol. XVI, London.
Beddard, F. E. (1911a). Contribution to the Anatomy and Systematic arrangement of the Cestoidea.
I. On some Mammalian Cestoidea. Proc. Zool. Soc ., London.
- (1911b). Contribution to the Anatomy and Systematic arrangement of the Cestoidea.
II. On two new Genera of Cestodes from Mammals. Proc. Zool. Soc., London.
- (1912). Contributions to the Anatomy and Systematic arrangement of the Cestoidea.
IV. On a species of Inermicapsifer from the Hyrax, and on the genera Zscbokkedla, Tbysano-
taenia and Hyracotaenia. Proc. Zool. Soc., London.
Meggttt, F. J. (1921). On a New Cestode from the Pouched Rat, Cricetomysgambianum. Parasitology,
Vol. XIII, No. 3, Cambridge.
4S7
MOSQUITOES AND OTHER BLOOD¬
SUCKING ARTHROPODS OF THE
UPPER SHIRI RIVER, NYASALAND
BY
Dr. J. B. DAVEY
AND
Professor R. NEWSTEAD
(Received for publication 23 November , 1921)
This paper deals with the mosquitoes and other sanguivorous
Arthropods observed in a relatively small area on the banks of the
Shiri River, a little south of Lake Malombe, in the Nyasaland
Protectorate, British Central Africa. Our captures were made
during the dry season, from July to the beginning of November, in
the year 1911. No special effort was made by us to collect such
material, and no search for the breeding-places of mosquitoes was
undertaken.
Our camp occupied a position about 200 yards from the river,
the intervening space being comparatively free from ‘ bush 9 or other
forms of vegetation. Hereabouts the river banks were low, and
partly or wholly submerged during the rains. Nearby the banks
were elevated and clothed with tall forest trees, and beneath
them there was a dense and almost impenetrable undergrowth.
Two-thirds of the river was rendered impassable by the ‘ sudd/ which
was composed of a dense floating platform of aquatic plants, serving
as a retreat for many species of birds, the crocodile and the
hippopotamus. The sudd consisted largely of grasses, and in places
extensive colonies of papyrus; the fringe being composed chiefly of
the beautiful ‘water caltrops' ( Trapa bispinosa ) and the equally
troublesome ‘ cabbage * or ‘ duckweed ' ( Pistia stratiotes ).
Mosquitoes (Culicidae).
Anopheles {Myzorhynchus) mauritianus , Grandpre. This fine
Anopheline was by no means common in our camp, as in all only ten
examples were captured ; nearly all of these came into our dining-hut
458
while we were sitting at the table after dark. It is just possible that
they may have been attracted by the artificial light; but of this we
could not be certain. It did not seek shelter in the tents, as in the
case of Anopheles funestus , and, therefore, does not appear to be a
strictly ‘ domestic* species. With one exception, the specimens were
of the dark form such as has been recorded from south of the
Zambesi, and were characterized by the absence of spots either upon
the costa or the fringe; the palpi presented three very narrow bands,
the tips were in most cases black. The dates of capture were: —
July, 1911; 1st August, 1911; 3rd August, 1911; 16th September,
1911; 18th September, 1911; 21st September, 1911.
Anopheles ( Cellia ) pharoensts , Theobald. One specimen only.
This was taken in the dark-room tent, 3rd August, 1911.
Anopheles ( Pyretophorus ) costalis, Loew. Only one example of
this mosquito was captured, during the month of August.
Anopheles ( Myzomyia ) funestus , Giles. This malaria-carrying
mosquito was the most abundant of all the Anophelines observed by
us. On July 31st, we decided to fix our permanent camp on the
banks of the Shiri, opposite Matutas village, but our tents became
so badly infested with this mosquito that we decided to abandon
the site, selecting a more open spot a short distance away. Our
first camping ground in this locality was surrounded by tall and,
for the most part, densely foliaged trees, and was distant from the
river about 80 yards. The permanent site of the camp was
200 yards from the water in practically open country. We found,
however, that this Anopheline was equally abundant in both places:
on August 2nd, one hundred and eleven specimens of this species
were counted in one of the tents, and there were certainly an equal
number present in the other tent. The temperature on this occasion at
3 a.m. was 46° F., at midday 86° F. in the shade, and at 8.30 p.m.
55 0 F. On August 9th, one hundred and sixty-two specimens of
this species were captured in one of the tents. These consisted
of thirty 3 3 > twelve unfed 9 9 and one hundred and twenty
9 9 which contained blood. This total does not, however, represent
the actual number present in the tent, as large numbers escaped and
many remained uncaptured, so that there were probably half as
many again as the total captures, and there was certainly an equal
number present in the other tent. This was the only occasion on
459
which we attempted to catch as many individuals as possible with
the view of ascertaining the ratio of sexes, the proportion of fed and
unfed females, and the approximate number present in the tent.
Further captures were made on August 14th and 31st, when
twenty-three 9 9 and three 3 3 * sixty-eight 9 9 and eleven 3 3
were captured respectively. We consider that the figures given for
August 9th may be taken as representing the approximate numbers
present daily in either tent at that period; but we noticed even
larger numbers on several occasions. During this time the tents
were opened daily at each end, and as many mosquitoes as possible
driven out, but as already stated we found no diminution in the
numbers. The largest numbers observed were always in the darker
portions of the tent, such as the angles of the roof and sides, between
the boxes and on various garments. We also observed that they
settled freely upon freshly skinned birds, especially guinea-fowls.
We also observed on many occasions large numbers flying into the
tents in the early morning between dawn and sunrise, the chief point
of entrance being at the upper portion of the opening, through
which they passed in more or less continuous flight until the rising
sun put an end to their movements. Our native employees, about
fifty in number, slept in the open about 15 to 25 yards from our
tents. The nearest native village being on the opposite side of the
river and about half a mile distant, we came to the conclusion that
the principal food supply of these mosquitoes was obtained from the
natives in our camp.
Mansonioides unifortnis , Theobald. It was by far the most
abundant of all the mosquitoes, and also the most vicious and
persistent biter. It attacked one at any time of the day or night,
but was most troublesome shortly after sunset. It simply swarmed
along the river and its immediate vicinity, but occurred also in the
bush in places far remote from water. It was not, however,
4 domestic * in its habits, though a few specimens were taken in our
tents during the day. It was equally obnoxious on the lower Shiri;
but almost entirely disappeared in the Zambesi where the margin of
the river was free from aquatic plants. It is highly probable,
therefore, that it breeds chiefly in those portions of the Shiri where
. the 4 sudd 1 is extensive, and where rootlets and stems of aquatic
plants suitable for the attachment of the larvae abound.
Culex- tigripes, Graitdpre. One feihale only was taken in one
of the tents in our camp during the month of August.
Ingsatnia ( Mifnomyta ) uniformis , Theobald. One female of this
rather rare mosquito was taken at night in our camp on August i ith,
1911. The beautiful pale blue reflections on the thorax were very
marked in this example.
Eiorleptiomyia mediolineata, Theobald. One female was taken
at the camp.
Taeniorhynchus aurites , Theobald. Examples of this species
were taken in the camp, the date for which is now lost.
PSYCHODIDAE.
Phlebolomus minutus var. africanus , Newst. Three examples
of this species were captured during the daytime, while at rest inside
the tents; and one at artificial light, at 7.30 p.m. Other specimens,
presumably of the same species, were also seen inside the tents at
various times generally resting upon the canvas roof. On two
occasions late at night examples attempted to bite one of us while
under the mosquito net; the familiar high-pitched note, somewhat
resembling that produced by a mosquito but much fainter, was heard
distinctly as the insects hovered round the face of its would-be
victim; on the first occasion we were sleeping under canvas; the
second time in our large mud-house.
The first example was taken in the middle of July; the others
were seen during August, and the last one towards the end of
September. It is evident, therefore, that this insect occurs in
Nyasaland during the dry season as it does also in Malta, and
possibly also in other parts of Africa.
TABANIDAE.
Tabanus taeniola var. varia/us, Walker, occurred very sparingly
towards the end of August, four specimens only being seen and
captured. 1'hese were all females. One was captured inside one
of our tents, one was caught on a dead hippopotamus, and two were
attracted by the fresh, moist mud on the walls of our hut.
Tabanus africanus , Gray. Two females of this handsome species
were also attracted by the fresh mud which covered the walls of the
hut. None was seen after the moisture had evaporated from the
mud.
461
PUPIPARA.
Echestypus sepiaceus , Speiser. A number of specimens were
taken from a young Kudu bull which was killed five miles west of
our camp on September 14th, 1911.
Olfeista ardeae , Macq. Two examples of this Hippoboscid
were taken from a freshly killed Goliath heron ( Ardea goliath ),
August 22nd, 1911. It is an extremely active insect, and the
specimens were caught with difficulty.
Hemiptera (Family Capsidae).
Trigonotylus brevipes> Jak. This minute green bug was first
observed in our camp an hour after sunset, oh August 10th, 1911,
when it bit one of us severely on the back of the hand; the proboscis
being driven so firmly into the skin as to prevent its immediate
escape, so that one was enabled to examine it carefully with a
pocket-lens. Subsequently this insect occurred in large numbers at
irregular intervals during August and September, always apparently
attracted by artificial light, and always annoying by its persistent
efforts to bite. It cannot, however, be considered a true blood¬
sucking insect, and we believe that its bites are made out of mere
curiosity rather than to obtain blood. In life, it is of an almost
uniform grass-green colour, with a narrow, pale streak below the
costa of the elytra, antennae dull crimson-red, basal portion of
terminal segment with an indistinct greyish ring; eyes black;
abdomen vivid green; legs slightly paler, especially the hind femora;
terminal segments of tarsi black.
Ticks. Ixodidae.
Though a number of animals were examined during our stay in
the Shiri Valley, comparatively few of them were found infested by
ticks. The following is a list of all the species observed.
Rhipicephalus neavei , Warburton. One male and four females
from a Kudu, five miles west of camp, 1911, many specimens
from a buffalo, 3rd August, 1911; two males and two females off
Nswala or Mpala antelope, Upper Shiri Valley, 8th August, 1911;
one female off an eland, Upper Shiri Valley, 20th August, 1911.
462
Rhtpicephalus falcatus , Neumann. Off buffalo, 3rd August,
I9n.
Rhipicephalus sintus , Neumann. Three males from buffalo,
Upper Shiri Valley, 3rd August, 1911.
Rhipicephalus maculatus i Neumann. One male from buffalo,
Upper Shiri Valley, 3rd August, 1911.
Hyalomma aegyptium , Linn. Off buffalo, Upper Shiri Valley,
3rd August, 1911.
463
BREEDING PLACES OF ANOPHELINE
MOSQUITOES IN FREETOWN, SIERRA
LEONE
BY
B. BLACKLOCK
(Received for publication 29 November , 1921)
Plates XXVII to XXXI and Three Maps
Stephens and Christophers (1900) stated ‘ in Freetown we found
that during the dry season the streams were the main source of
Anopheles / Daniels (1901), in a letter to Sir Ronald Ross bearing
on the anti-mosquito operations then proceeding in Freetown,
expressed the opinion that in the dry season, while the surface
collections of .water would have disappeared, new breeding-places
would arise, * mainly the streams, small and large, which remain,
possibly some of the other wells and artificial collections of water in
tubs, etc.’ Boyce, Evans and Clarke (1905) mention ‘the great
disadvantage which attaches to the streams in the dry season, viz.,
that they constitute in Freetown, at that period, the chief sources
of Anopheles supply/
With the object of ascertaining whether the streams referred to
above, which traverse Freetown, still act at the present day as
breeding-places of anopheline mosquitoes, I took the opportunity of
examining them at the end of the dry season of 1921. The search
for larvae was carried out in the month of May, at which time the
stream beds contained comparatively small amounts of water. The
plan adopted in making the survey was to start at the mouth of the
stream at the point of entry into the sea and to work up stream
through the town until the places of origin of the streams was
reached in the high ground at the foot of the hills behind the town;
the various tributaries were followed in turn.
The results obtained showed that in each stream in which larvae
were found, breeding-places were present in largest numbers at the
4 6 +
lower end of the stream; as one proceeded upwards through the
town the. breeding-places and the numbers found became scanty,
while on emerging from above the town breeding places were again
found, but still in small numbers in comparison with those at the
lower end of the streams.
The reasons for this distribution are doubtless numerous, but
some of them are clearly connected with the conformation of the
ground. The streams, in some cases, by the time they reach a point
several hundred yards from the sea have produced wide, steep-sided
gorges through which they pursue a tortuous, irregular course; the
numerous sheltered bends, overgrown with grass and weed, afford
excellent breeding-grounds and shelter for the larvae. The process of
erosion in the wet season appears to have more than compensated for
the additional volume of water to be carried, so that the advantage
of flushing is to a certain extent lost in the lower portions of the
streams. In the town proper the streams are passing through a rocky
formation, with the result that there is slower erosion; this and the
great amount of canalisation of tributaries which has been carried
out in recent years permit of more complete washing out of the
stream bed in its passage through the town. Above the town the
prevalence of larval breeding-places was to be anticipated from the
more diffuse nature of the water courses, their extensive area, and
the plentiful vegetation through which the water slowly percolates
before forming definite streams. The conformation of the ground,
while it favours the presence of .Anopheles by providing suitable
breeding-places, acts also in their favour, especially in the gorges
mentioned above, by rendering it difficult to obtain access to these
breeding-places for the purpose of detecting them, and still more by
rendering the application of the measures necessary for their eradica¬
tion very laborious.
The maps appended show (Nos. I and II) the places in which
anopheline larvae were discovered and the manner of their distribu¬
tion in 1921 and 1900. It was observed that the residual breeding-
places in the streams at the end of the dry season are of two kinds.
The first is the edge of the winding and eroded bed of the stream
just before its entrance to the sea, the second is the shallow water,
well protected by vegetation and extending over a large surface,
which is found at the places of origin of the streams. These residual
Map i. # A. costalis lame or pupae found 1921.
Map 2. Anopheles larvae found 1900.
4 66
breeding-places doubtless contribute very materially to the spread
of Anopheles when the rains commence, and other breeding sites
become available. Some of the more interesting breeding-places
found are seen in photographs i to 5.
SPECIES OF ANOPHELES FOUND *
Larvae and pupae collected from the various breeding-places
were taken to the laboratory and allowed to develop ; anophelines
which emerged belonged in all cases to the species * 4 . costalis. This
appears at the present time to be by far the commonest anopheline
breeding at the end of the dry season in Freetown. It is deserving
of note that in the early days of mosquito investigation in Freetown
this species predominated at the end of the wet season also.
Daniels (1901) mentions Anopheles costalis as a common mosquito
in Freetown, and further states that * A. funestus was found near
but not in Freetown/ Ross, Annett and Austin (1902) stated that
while of over two hundred anophelines obtained from Wilberforce
Barracks all were A. costalis , they obtained from Dr. Berkeley at
Kissy both this species and also A . funestus , the latter 1 by far the
more numerous/ They add that A. funestus was restricted entirely
to the eastern part of the town, and that they never found a larva
or an adult of this species west of Government House. It is not
stated, however, how near to this dividing line A . funestus existed
on the eiist side at that time. Thus, while there is no evidence
from the foregoing and the present survey that the proportion of
A. funestus to . 4 . costalis has undergone a change during this period
of years, there is considerable evidence to show that A. costalis was
then, and is to-day, the commonest anopheline in Freetown.
Owing to lack of time, little could be done in the way of dissections
of adults, either caught wild or experimentally fed on malaria
carriers; this will be reserved for a further date, when it will be of
great interest to discover whether there exist to-day any rates of
mosquito infection comparable with the Wilberforce Barracks
figures of Ross and collaborators in 1902, namely, twenty-seven
infected Anopheles costalis in one hundred and nine captured females
dissected.
Annals Prop. Med. & Parasitol., J’ol. XV
PLATE XXVIII
Photograph No. 2
Anopheline breeding place, above the town. The water is sluggish and almost concealed by
vegetation. Numerous A. costal is larvae found to left of man.
C. Tifiling Co., Ltd., Imp.
+$7
It may be said then that the results obtained in 1921 at the end
of the dry season were such as to confirm the predictions of twenty
years ago, and in view of the great amount of careful and pains¬
taking work which has been steadily carried out in the interval with
a view to abating mosquito breeding, it is clear that in these
streams we have a problem which will require a little more detailed
investigation.
METHODS OF DEALING WITH THE STREAMS
Previous suggestions as to how such an obvious source of
Anopheles could be dealt with have been of various kinds. Some
of these may be mentioned;
Daniels (1903) wrote: ‘Two possible methods which are most
obvious are the formation of a central channel in the bed of the
stream, with larger collections of water in sufficient numbers of
places for drinking purposes, and, lower down the stream, other
places for washing, etc. The second, which might be cheaper but
certainly less effective, would be to dam up the streams so as to
obtain a sufficient head of water to flush out the whole channel at
intervals.*
Boyce, Evans and Clarke (1905) summarised their views on. this
matter as follows : —
(1) Reconstruction of the bed of streams.
(2) Diversion of water for flushing of town drains in dry
weather.
(3) Construction of dams in streams and flushing of beds at
intervals.
It will be seen that schemes one and three here correspond to the
two suggested by Daniels above. The town drains referred to, in
number two, are the surface drains in the streets.
(1) Reconstruction of the bed of streams.
If one follows such a stream as Nicol*s Brook from its origin above
Foulah town to the point where it enters the sea, it is realised that
the operations involved in this scheme would be of great magnitude.
The process of erosion has been so irregular in its action that the
water does not flow on a simple slope from above down to the sea
468
level. It forms at places considerable falls, and also has cut very
large cavities in the bottom of the stream bed. Laterally also it
extends in different places to very different dimensions. In order to
rectify the levels and build a channel capable of carrying all the
rainy season water, it appears that a very large expenditure would
be involved. If when this expenditure had been made the streams
could then be regarded as safe, it might be a plan worthy of
consideration : but it appears probable that constant attention would
be required to ensure that the lateral tributaries discharged properly
into the central channel and that pools did not form outside this
channel itself. This would involve one of two things. Either the
channel would have to be so levelled and sloped at the sides that no
pools could possibly form—a vast operation—or else such pools
would have to be treated regularly with larvicide, as at present.
The other streams would require similar measures.
(2) Diversion of water for Hushing of town drains in dry weather .
This % scheme would, it appears to me, fail to be effective in
preventing anopheline breeding in the stream beds, for two reasons.
One is that water oozing out of springs between the strata in> the
bed of the stream would still form pools suitable for breeding
anophelines. The other is that many of the drains referred to
discharge into these streams and would thus carry the water back
to the bed of the stream after having flushed out the town street
drains.
(3) Construction of dams in streams and flushing of beds at
intervals .
This scheme assumes that the bed of the stream is capable of
being effectually flushed, but the observations which were made
above in connection with the conformation of the streams go to
prove that Nature has already by the process of excavation and
erosion effectually prevented such a flushing action being successful.
Further, it cannot be said that even a large volume of water passing
occasionally down a river bed which is a honeycomb of pools will
ensure the sweeping out of anopheline larvae from these pools.
This scheme, therefore, does not appear to guarantee success.
Annals Prop. Med. & Parasitol ., Vol. XV
PLATE XXIX
Photograph No. 3
Anophclinc breeding place, in stream bed in the town,
bank to left of where the woman is standing.
A. costal is larvae found near
C. Ti tiling <0? Co., Ltd., Imp.
469
SEGREGATION METHOD
Stephens and Christophers were among the pioneers of the
segregation method of dealing with the malaria problem, and there
i$ ample evidence of the efficacy of this method where it is vigorously
and thoroughly carried out. It is chiefly of value where we are
dealing with a small non-immune community which has to exist side
by side with an overwhelming preponderance of immunes and semi-
immunes. It is based on the fact that there always has been, and
always is, a large proportion of infected children among the native
population. If we believe such conditions will always remain in
the future, it is clear that the readiest and most inexpensive way of
dealing with such a small white community will be by strict
segregation; in this way a great degree of safety from infection can
reasonably be insured. In places which are being newly developed
this method can be used advantageously, as also in places where
the white population is small and circumstances favour their settle¬
ment away from the native. But in old established towns it is not
by any means an easy method to apply, more especially where the
white population is growing, and likely to grow greater year by year.
It is also a method which leaves the natives to themselves and frankly
regards them as so infected that no immediate and inexpensive
mode of dealing with this infection is available; to this extent it is
a method which is a tacit acknowledgment of defeat. Segregation
occupies in respect to the malaria problem much the same place
from the point of view of medical prophylaxis as evacuation of fly
areas occupies in respect to the trypanosomiasis problem. Each may
be excellent as a temporary expedient, but neither can be regarded
as a final and satisfactory method of dealing with these problems.
The bearing of segregation on the question of how to deal with
the streams of Freetown is clear when we consider the results
obtained by Stephens and Christophers in their investigation on the
anopheline content of these streams and the native houses situated
dose to them. I have obtained the permission of these authors to
reproduce a spot plan (Map III) which they made in 1900, showing
the distribution of adult anopheles in houses near such a stream.
It will be observed that ‘houses in which anopheles are present in
enormous numbers * are found immediately beside the stream, that
470
‘ houses in which, search in early morning reveals a few anopheles *
extend back from the stream a short distance, while as we recede
further from the stream we reach ‘ houses in which anopheles cannot
be found/ The conditions depicted here would, I feel no hesitation
in saying, reproduce themselves automatically immediately any
considerable relaxation of the present sustained effort on the part
Houses ir\ which anooheles a^e present Wnprfr\ous nun\\>^rs
■ Houses ti\which search mcarl^ morrvrvo reveals afew anopheles
O Houses try which anopheles cannot he found
Map IH.
of the sanitary authorities of Freetown took place. In Freetown it
does not appear to me practicable now to adopt a method of rigid
segregation, and moreover, even if it were practicable, it would still
be advisable to deal with the problem of the streams with a view
to ameliorating the condition of the native.
Annals Trop. Med. & Parasitol. , Vol. XV
PLATE XXX
Photograph No. 4
Anophelinc breeding place, in the town. A few A. costalis larvae were found at points
along the edge from where the man is standing ; water fairly rapid.
C, Tinting & Co., Ltd,, Imp,
47 1
IS A PERMANENTLY EFFECTIVE SCHEME POSSIBLE?
The only permanent scheme which will be effective in dealing with
the stream beds appears to be one which obliterates them entirely.
Whether any such scheme can be devised is a matter for engineers to
determine. It might involve the construction of a canal or canals
capable of carrying the water to east and west from above the town
and reclamation of the present stream beds. The fact that the
outlay required, if the scheme were practicable, would be very large
indeed does not enter at present into the discussion. Whether the
effective method will prove too costly need and can only be decided
after considerable investigation over a period as to whether an
effective scheme is practicable from the engineering point of view.
It deserves investigation, however, even if apparently excluded on
account of prohibitive cost, for we are not justified in assuming that
an effective method which would be too expensive at present must
necessarily prove so in future years.
REFERENCES
Boyce, R. Evans, A., and Clarke, H. H. (1905). Report on Sanitation in Freetown. Liv. Scb.
Trop. Med. Memoir. Vol. XIV, pp. 32 and 33.
Daniels, C. W. (1901). Liv. Scb. Trop. Med. Memoir V. Pt. I. Appendix, p. 15.
Ross, R. Annztt, H. E., and Austen, E. E. (1902). Report of the Malaria Expedition. l>iv.
Scb. Trop. Med. Memoir II, p. 10. »
Stephens, J. W. W., and Christophers, S. R. (1900). ‘ The Native as Agent in the Malarial
Infection of Europeans.* Reports to the Malaria Committee of tbe Royal Society , p. 6.
Annals Trop. Med . & Parasitol ., Vol. XV
PLATE XXXI
Photograph No. 5
Anophclinc breeding place, in the town. Behind the rock in front of which the man is
standing is the pool in which little vegetation was present ; a few A. costalis larvae.
C, Tifiling & Co., Ltd., Imp.
473
NOTES ON AN APPARATUS FOR THE IN¬
DIVIDUAL BREEDING OF MOSQUITOES
/ •
BY
B. BLACKLOCK
t
(Received for publication 29 "November , 1921)
The apparatus here described is designed to facilitate the task
of breeding out mosquitoes individually from any stage in which
they happen to be at the time when they are collected, and in this
way obtaining a record of the association of the various stages. It
is a development of .the breeding apparatus which Carter and
Blacklock (1920) described, and its small size makes it easily
portable and convenient for use in the tropics. By means of it, with
a little trouble, a complete record of the stage originally obtained,
the a.dult stage, and the intervening stages is possible. The parts
of which the apparatus consists are glass capsules of suitable size,
embedded in a wooden base, glass cylinders of which one end is
covered with mosquito netting, and glass tubes for the collection of
the casts at the various stages of development. It will be simplest
to give the dimensions of the model actually in use; this has proved,
in so far as mosquitoes are concerned, satisfactory, the mortality,
whether of larvae, pupae or adults, being comparatively small.
Modifications could easily be introduced which would render the
apparatus suitable for observing the development of other insects in
their aquatic stages.
The capsules. These are made from glass tubing, and are of the
following dimensions:—Internal diameter, 11 mm.; internal height,
18 mm. They are of stout glass, about 1 mm. thick, rounded at the
bottom like a test tube (fig. 1), and they fit into holes drilled in the
Fig. i.
wooden base at measured intervals. The depth of the hole bored is
such that when a capsule is placed in it the edge of the capsule lies
474
flush with the surface of the board, and the width of the holes is
such that it is possible with fine bladed forceps to lift the capsule
out of the board. A small ring of plasticine can be used to fit the
junction of the holes that are too large for the capsules.
The board . The wood used in making the board is hard and
well seasoned so that it will not warp. The length is 30 cm., the
width 11 cm., the depth 3 cm.; the holes are bored in three rows,
the distance between the holes from each other in any row or between
the rows being 2 cm. (fig. 2). In this manner it is possible to have
FlG. 2.
in each board twenty-seven capsules arranged in three rows of nine
capsules each. In boring the holes, the first row is bored near the
edge of the wood, leaving a rim of only about 0 5 cm. in front of
them; the object of this arrangement will be explained later when
describing the manner of using the apparatus.
The cylinders . These are made of glass, of similar thickness
to that used for the capsules. The dimensions are:—Internal
diameter, 15 mm.; height, 25 mm. It is essential that one end at
least of the cylinder should be so cut that it will stand upright and
make fair contact all round its edge with the board on which it
stands, if one end is irregular, it should be used as the upper end;
this end is in all cases covered with a small piece of mosquito netting,
kept in position by a piece of string or two thin rubber bands (fig. 3).
Fig. 3.
475
As rubber is apt to deteriorate in the tropics and a band may break,
it is necessary when using such bands to use two to ensure safety;
string, however, answers the purpose quite satisfactorily, merely
taking a little more time to fix on and take off. Fig. 4 gives a
section of the cylinders in position covering the capsule embedded in
the board.
Tubes for casts . These are made from glass tubing of an internal
diameter of not less than 5 mm. and about 6 cm. in length, drawn
to a blunt point at one end (fig. 5). When the larval and pupal
t .
Fig. 5.
pelts have been placed in them they can be corked and waxed, or
better, if so long as to permit of it, they can be drawn out in a small
flame and sealed.
Method of use. The stages collected in the field, and from which
it is desired to breed adults, are placed in a dish of such a size as to
render it easy to ensure the isolation of individuals, and one. by one
these are transferred to the capsules, each into one capsule; this is
filled with water and replaced in its position in the board, as
evaporation occurs the fluid lost must be replaced daily to keep the
water at the surface level. This is especially necessary in the case
of anopheline larvae. The capsule is marked by a number written
+76
on the board in front of it in pencil, the same number being entered
in the book in serial order. The front of the board is that margin
on which the narrower rim is left; this serves to identify the front,
and is useful in practice also by leaving standing room at the back
of each capsule for the accommodation of the cylinders when they
are removed from their capsules. The capsule having been filled with
water and replaced, a small portion of dried, powdered cockroach,
or other suitable *food material, is deposited on the surface of the
water with a needle. Each day the capsule is examined at intervals,
and when a cast skin is observed it is picked out with a needle
or fine camel-hair brush and transferred to one of the tubes which
have been described above, this being filled half full of preserving
fluid, or the whole contents of the tube, including the larva, may be
pipetted out with a wide bore pipette into a small porcelain dish
and the cast skin recovered, the larva being replaced. Transfer by
means of a pipette is apt to lead to difficulties owing to the fact that
whereas the larval pelts usually fall to the bottom, the cast pupal
skins usually float and may become stranded, on the side of the
pipette used. The date of the stage is noted in the record, the tube
is numbered with the same number as the capsule and is retained for
all material which is derived from that capsule. As soon as the
pupal stage is reached one of the cylinders is placed in position over
the capsule, and when the adult emerges into the cylinder this may
be pushed carefully back off the capsule on to the board immediately
behind it, in order to give the adult time to dry and expand. At
the same time the pupal cast is removed and placed in its tube,
which is then corked and waxed or sealed over a flame. When the
adult is ready for pinning it is removed from the board by slipping
a strip of thin, stiff paper underneath the cylinder containing it,
chloroformed by sliding the cylinder on to a piece of paper moistened
with chloroform, while the upper end of the cylinder is covered by
the finger; when pinned it is labelled in the collecting box with the
same number as the capsule from which it was derived and the tube
containing its other stages. Before use again after one experiment
the capsule is carefully swabbed out with a pledget of cotton wool,
first with water, then with alcohol, and dried.
In this way it is possible to obtain in the majority of cases a very
complete record of the stages from the time the individual was
477
collected; the apparatus is of value, not only as a simple means of
associating stages and noting their duration in the laboratory, but
also for isolating mosquitoes as they emerge if required for experi¬
mental purposes.
Precautions in use . Care is required with reference to the
following points: —
(1) Proper entry in the record of the number given to the
capsule, pelt-tube and adult.
(2) That one individual only is introduced into the capsule;
this is especially necessary when dealing with young
larvae or eggs.
(3) That each pelt is removed as soon as it is shed.
(4) That the tube containing pelts is well sealed if intended to
be kept for future examination.
479
THE TREATMENT OF A CASE
OF RHODESIAN SLEEPING SICKNESS
BY THE PREPARATION KNOWN AS
‘BAYER 205’
BY
WARRINGTON YORKE
(Received for publication 6 December , 1921)
The patient (G. J.) contracted the disease in the Seringe district
of Nofth Eastern Rhodesia on the Congo Zambesi Watershed.
He was suddenly taken ill on 17th September, 1920, with severe
headache, pains in the back and neck, great prostration and a
temperature of 105° F.; during the first eight days of the illness the
temperature varied between io4 °F. and 106° F. Trypanosomes
were found in the blood on 27th September, and the patient was
thereupon sent to Broken Hill, where he arrived on 24th October,
and was treated by Dr. Wallace with tartar emetic, intravenous
injections (2*5 to 3 grains) being given every other day. A little
later intramuscular injections of antimony oxide (/ a grain) and
soamin (2 grains) were also given. Between 24th October and
26th December 3*28 gm. of tartaric emetic, o*i gm. of antimony
oxide and 2 47 gm. of soamin had been administered, with only a
moderate degree of benefit to the patient: trypanosomes, were
frequently found in the blood and there were several severe febrile
disturbances. Between 26th December and 15th January, 1921,
thirteen injections of stibenyl (in all 3 05 gm.) were given. These
injections were, however, badly borne, and as trypanosomes were
present in the blood on 16th January, the original treatment, with
tartar emetic, antimony oxide and soamin, was resumed. Things
improved somewhat, and on the 3rd February the patient was well
enough to leave for England. At Cape Town, and on the voyage,
he had further injections of tartar emetic.
When the patient reached Liverpool on 12th March, he was in
fairly good condition; trypanosomes were not present in his blood
480
(inoculated rats did not become infected), there was well-marked
autoagglutination of the erythrocytes, and the weight was 144 lbs.
Daily examinations of the blood were negative until 18th March,
when a few trypanosomes were found. Inoculation of rats showed
the parasite to be T. rhodesiense. Intravenous injections of tartar
emetic in doses of from 2 to 3 grains were then given on alternate
days. Neither these nor injections of atoxyl, however, seemed to
exert any influence on the course of the disease. Violent febrile
disturbances accompanied by severe headaches and great prostration,
and the appearance of trypanosomes in the peripheral blood in
considerable numbers, occurred with the utmost regularity every five
or six days, and the patient went steadily down hill.
In view of the fact that the infection appeared to be antimony
and arsenic resistant, I endeavoured to obtain from Professor
Mayer, of the Hamburg School of Tropical Medicine, a small
quantity of a preparation called ‘Bayer 205’, manufactured by the
firm of Friedr. Bayer & Co., Elberfeld. Although this drug—the
constitution of which was not disclosed on account of the state of
the Germany chemical industry—had not been used in any case of
human trypanosomiasis, it had been employed with remarkable
results in the treatment of experimental trypanosomiasis of animals,
records of which had been recently published by Haendel and
Joetten (1920) and by Mayer and Zeiss (1920).
Professor Mayer informed me that he was unable to comply with
my request to let me have a supply of the drug for trial, but stated
that if I sent the patient to Germany they would be glad to treat
him at the Hamburg School. This, unfortunately, resulted in some
loss of time, as I did not like the idea of sending the patient to
Germany to be treated with a proprietory article, the composition
of which was not disclosed and which could not be sent out of the
country. However, as notwithstanding the most rigorous treatment
the patient’s condition got steadily worse, I went critically through
the above mentioned papers again and was so impressed by the
results recorded, that I overcame my own scruples and those of the
patient, with the result that he left for Hamburg on 5th July, 1921.
During his eight and a half months illness the patient had received,
in addition to the other drugs mentioned, 22*7 gm. of tartar emetic,
and at the time he left for Hamburg his weight had fallen to
481
132 lbs., he was suffering from frequent violent febrile disturbances,
antimony and arsenic preparations were without any effect on the
course of the infection, and the condition seemed almost hopeless.
The following details of the case are taken from a recent paper
by Muhlens and Menk (1921), under whose care the patient was when
in Hamburg. On 9th July, two days after his arrival, he had a
rigor, accompanied by headache and fever, and trypanosomes were
found in the blood. Treatment was immediately commenced,
1 Bayer 20.5’ 0 5 gm. in 5 per cent, solution being given intra¬
venously. The injection was well tolerated; the fever subsided in
six hours instead of in the customary thirty-six hours, and trypano¬
somes had disappeared from the blood within sixteen hours. The
next day a second injection of 1 gm. in 10 per cent, solution was
given, and this was repeated on the following day: both injections
were well borne and the urine remained free from albumen or casts.
The blood was examined twice daily during the following seven
days, with negative results, and a further injection of 1 gm. was
then given. As a small quantity of albumen and occasional red
and white blood corpuscles appeared in the urine, no further treat¬
ment was administered. The patient improved remarkably, rapidly
regained his sense of well-being and put on weight. Almost daily
examinations of the blood were made, until 29th August, but with
negative results: there were no further febrile attacks or any other
untoward symptoms, and on 13th September the patient returned to
Liverpool.
When I saw him on 15th September, I was greatly impressed with
the remarkable improvement in his general condition: he had lost
the anxious and weary expression which had been so characteristic
for some months before he went to Germany, his weight had increased
to 145 lbs., the blood was negative and sub-inoculated rats did not
become infected, and the autoagglutination of the erythrocytes had
disappeared. The urine contained a trace of albumen, otherwise the
patient seemed perfectly well. The following day he left Liverpool
for his home in South Wales. On 5th November, 1921, he returned
to Liverpool for further examination. He appeared in excellent
health, and stated that he never felt better in his life aind had played
two rounds of golf daily without the slightest distress. The blood
was examined microscopically and also inoculated into three rats,
482
with negative results: there was no autoagglutination ,of the
erythrocytes or any other sign of the disease: the weight had
increased to 150 lbs. (patient's normal weight), the urine, however,
still contained traces of albumen. On nth November he sailed for
South Africa.
The above record is of peculiar interest, as it relates to the first
case of human trypanosomiasis treated with adequate doses* of
1 Bayer 205 \ Whilst recognising that hasty and premature claims
of success in the treatment of a disease like sleeping sickness are to
be deprecated, and that the further history of the patient alone can
decide whether a real therapia sterilisans magna has been effected,
the facts that the disease was of at least eight and a half months
standing at the time of treatment with ‘ Bayer 205 ’; that the patient
was extremely ill and had been getting steadily worse for months,
that the infection was completely resistant to the ordinary prepara¬
tions of antimony and arsenic; that since the administration of
1 Bayer 205 9 all symptoms of the disease have disappeared, the
general condition of the patient has improved enormously, his weight
has increased by 18 lbs., his blood has been free from parasties and
non infective to rats for an observation period of four months—these
facts, considered in the light of the usual history of cases suffering
from T. rhodesiense , warrant the hope that in * Bayer 205 ' we have
a drug of exceptional trypanocidal power.
REFERENCES
Haendel, and Joetten (1920). Berlin Klin . Week., Vol. LVII, p. 821.
Mayex, and Zeiss (1920). Arcb.f. ScbiJJs - 11. Trop.~Hyg. } Vol. XXIV, p. 257.
MUhlens, and Menk (1921). Miincb . Med, Wocb. y Vol. LXVIII, No. 46, p. 1488.
• M iliac us and Meiik (loc. cit.) give details ol the administration of the drug to a case
of gambiense infection in March, 1920, but only two small doses (0*2 gm.) were given at an
interval of a week. A relapse occurred and the patient was then treated with other drugs.
INDEX
INDEX
PAGE
Index of Authors . iii
General Index ./. iii
Index of Genera, Species and Varieties new to Science . xi
INDEX OF AUTHORS
PAGE
Adler, S.427, 433
Adler, S.; and Stephens, J. W. W.... 173
Adler, S.; and Yorke, W.:. 269
Allmand, D. 1
Barret, H. P... 113
Blacklock, B.59, 463,473
Blacklock, B.; and Carter, H. F. 73
Breinl, A. 49
Carter, H. F.; and Blacklock, B. 73
Carter, H. F.; Ingram, A. ; and
Macfie, J. W. S. 177
Davey, J. B.; and Newstead, R. 457
Evans, A. M. 445
Evans, A. M.; and Newstead, R....95, 287
Gordon, R. M.; and Young, C. J. ... 265
Hill, G. F.91, 93
I hie, J. E. W. 397
Ingram, A.; and Macfie, J. W. S. 53, 283,
313
Ingram, A.; Carter, H. F. ; and
Macfie, J. W. S. 177
Kennan, R. H. 245
Macfie, J. W. S.271, 377
PAGE
Macfie, J. W. S.; and Ingram, A. 53, 283,
313
Macfie, J. W. S.; Carter, H. F. ; and
Ingram, A. 177
MacGregor, M. E. 417
Maplestone, P. A.403, 407, 413
Maplestone, P. A.; and Southwell, T. 455
Newstead, R. ; and Davey, J. B. 457
Newstead, R.; and Evans, A. M....95, 287
Newstead, R. ; and Sinton, J. A. 103 4
Scott, H. H.117, 133, 149, 213, 227
Sinton, J. A. 107
Sinton, J. A.; and Newstead, R. 103
Southwell, T.161, 167, 169
Southwell, T.; and Maplestone, P. A. 455
Southwell, T.; and Yorke, W. 249
Stephens, J. W. W. 435
Stepheiis, J. W. W.; and Adler, S.... 173
Yorke, W. 479
Yorke, W.; and Adler, S. 269
Yorke, W.; and Southwell, T. 249
Young, C. J. 301
Young, C. J.; and Gordon, R. M.... 265
GENERAL INDEX
PAGE
Acanthotaenia gallardi in a new host... 404
Adler, S. Note on bismuth as a try-
panocide . 433
Adler, S. The trypanocidal effect of
phenylglycine amido arsenate of
sodium on T. brucei in rats and T.
rhodtsiense in mice . 427
Adler, S., and Stephens, J. W. W. A
case of suspected leprosy. 173
Adler, S., and Yorke, W. Note on a
case of leprosy. 269
Aedcs jcapularis .445, 446
| PAGE
Aedes serratus .* 445
„ (JStegomyta) fascia ta . 447
„ trivitiatus . 446
Aeschnidae destructive to S . calopus ... 302
| African rats, Cestodes from . 167
Allmand, D. Liverpool School of
Tropical Medicine: Scientific
Record . 1
I Alveatum-catinatum group of Cylicos -
tomum . 398
I Amoebiasis research by the Liverpool
School . 34
iii
PAGE
Aneurysms, multiple, in a child. 245
Angularia australis , sp.n. 407
„ genus, Diagnosis of .. 412
Ankylostoma in symptomless carriers.... 123
Anopheles , Breeding places of, in Free¬
town, Sierra Leone . 463
found on a Venezuelan oil¬
field . 437
„ albimanus . 446
„ „ morphological
characters ... 447
„ „ var. tarsimacula-
tus.,, .. 446
» argyrotarsus . 439, 445, 446
„ „ morphological
characters.... 447
„ bifurcatus in the Isle of
Man .73, 76
„ „ Ova of. 421
„ „ opposition. 422
„ costalis f commonest species
at Freetown,
Sierra Leone... 466
„ „ in Nyasaland. 458
„ funestus in Nyasaland . 458
„ maculipennis in the Isle of
Man .74, 82
» » Ova of .. 419
„ „ opposition ... 422
„ mauritianus in Nyasaland... 457
„ pharoensis in Nyasaland. 458
„ plumbeus in the Isle of
Man.75, 85
» „ Ova of. 421
„ „ opposition. 423
„ pseudopunctipennis in Vene¬
zuela . 446
Anoplocephala magna , var. pediculata... 256
„ perfoliata y Anatomy of 257
„ rhodesiensis , nom. nov.,
Anatomy of. 249
Antimony in treatment of Oriental sore 107
Apparatus for the individual breeding
of mosquitoes . 473
Arneth count in hookworm-infected
children in North Queensland . 49
Ascaris in symptomless carriers. 123
Atrichopogon africanum , sp.n., female 334
„ chrysosphaerotum , sp.n.,
female. 337
„ elektrophaeum , sp.n.,
female. 335
PACE
Atrichopogon homoium , sp.n., female ... 338
„ ptrfvscum, sp.n., female... 337
Australia, Arneth count in children in 49
„ Muse a domes tica as a 1 bush-
fly ’ in. 93
„ Unusual breeding-places of
Stegomyia fas data in. 91
Australian aborigines, Ulcerative granu¬
loma among. 413
„ cestodes .403, 407
Bacillus tuberculosis , Hereditary trans¬
mission of. 216
Balantidium coli> Incidence of, in
symptomless carriers.120, 128
Barret, H. .P. A method for the
cultivation of blastocystis . 113
Belgian Congo, New tsetse-flies
from . 95
Beri-beri research by the Liverpool
School . 35
Bezzia foyi , sp.n., female . 361
Bismuth as a trypanodde . 433
Blacklock, B. Breeding-places of ano-
pheline mosquitoes in Freetown,
Sierra Leone . 463
Blacklock, B. Notes on an apparatus
for the individual breeding of mos¬
quitoes . 473
Blacklock, B. Notes on a case of
indigenous infection with P. fold -
parum . 59
Blacklock, B., and Carter, H. F.
Observations on mosquitoes in the
Isle of Man. 73
Blackwater research by the Liverpool
School . 14
Blastocystis, a distinct zoological genus 116
„ Method for cultivation of 115
,1 Nature of . 113
Blood-sucking arthropods of the Upper
Shiri River, Nyasaland. 457
Breeding of mosquitoes, Apparatus for 473
„ places of anophehne mosqui¬
toes in Freetown, Sierra Leone. 463
Breinl, A. On the ‘ Arneth Count *
in hookworm-infected white chil¬
dren in North Queensland . 49
Brevicapsulatum group of Cylicostomum 401
Bronchomoniliasis complicating pul¬
monary tuberculosis in a native of
the Gold Coast .53, 285
Calicatum group of Cylicostomum . 399
IV
PAGE
PAGE
Carter, H. F., and Biacklock, B.
Observations on mosquitoes in the
Isle of Man. 73
Carter, H. F., Ingram, A., and Macfie.
Observations on the Ceratopogonine
midges of the Gold Coast, with
descriptions of new species. 177
Cellia argyrotarsis in Venezuela. 439
CeratophyUus jasciatus on rats at Liver¬
pool . 292
„ londiniensis on rats at
Liverpool . 294
Ceratopogoninae of the Gold Coast, 177, 313
Cestode, new, from a cormorant . 169
Cestodes from African rats . 167
,, Australian .403, 407
„ from Indian poultry. 161
„ List of, occurring in fowls 161
„ in new hosts . 403
Cbilomastix mesnili , Incidence of, in
Cidicoides inomatipennis , var. rutilus ,
v.n. 326
„ neaveiy pupa . 320
„ nigeriae , sp.n. 325
„ similisy larva . 322
„ „ pupa . 32 1
Cylicostomum , on the genus. 397
Alveatum-catinatum
group of . 398
Brevicapsulatum group of 401
Calicatum group of ... 399
Euproctus-bicoronatum
group of . 400
Montgomeryi group of... 401
Radiatum-elongatum
group of . 399
Tetracantbum-coronatum
group of . 397
labia turn, var. digitatum ,
var. nov. 401
symptomless carriers.120, 128 ! Dasyhelea genus, morphology of adult 178
Cittotaenia tackyglossi
4°3
larva 183
Cormorant, New cestode from a . 169
pupa 182
Cotugnia digonopora in Indian poultry 163
99
Synoptic table of new
Crescents, ‘ Immunizing 9 process
re-
species of.
211
suiting in .
71
boothiy sp.n., male.
331
„ Significance of .
70
jlava , sp.n., adult .
196
„ Time of appearance of.
68
„ » larva .
199
Cropper and Row’s concentration
99
„ „ pupa .
198
method of examining stools ....
118
99
flaviformisy sp.n., adult ...
201
Ctenoeephalis cants on a rat at Liver-
99
„ „ pupa.
203
pool .
294
99
fusca y sp.n., adult .
204
Ctenoptbalmus agyrtes not found on rats
99
« „ pupa .
205
at Liverpool.
295
99
Jusciformisy sp.n., female ...
209
Culex chrysotborax .
445
99
» » larva.
210
,, coronator .
44 6
99
„ „ pupa..
210
„ nigripalpus .
446
99
fuscipleurisy sp.n., female ...
200
„ pipiens in the Isle of Man ...
...75,88
99
» „ larva.
327
„ quinquefasciatus .
44 6 > 447
99
,4 „ pupa.
327
» tigripes .
46O
99
fusciscutellatay sp.n., adult...
187
„ virgultus .
446
99
inconspictiosay sp.n., adult....
i 9 i
Culicella fumipennis in the Isle
of
99
,, „ larva ...
194
Man .
...76, 90
99
„ „ pupa....
193
,, . morsitans in the Isle of Man, 76,90
99
luteoscutellatay sp.n., female
191
Culicidae collected in Nyasaland...
457
99
nigeriae, sp.n.
329
„ „ Venezuela .
445
nigricans , sp.n., male .
„ „ female.
194
Culicoides austeni , larva .
3*5
328
» „ pupa .
3 i 5
99
nigrofuscay sp.n., male .
207
„ corsoni , sp.n., male.
324
99
» pupa .
208
„ distinctipennis, pupa ....
316
99
pallidihaltery sp.n., adult ...
184
„ eriodendroni , male .
3 i 7
99
„ „ larva ...
186
„ grahamiy pupa .
318
99
„ „ pupa ...
186
v
PACE |
Dasyhelea retorta , sp.n., female. 333
„ similis , sp.n., adult . 189 j
Davainea genus, Fiihrmann’s revision 1
of. 162 |
„ celebensis from an African rat 167
„ cesticillus in Indian poultry 163 |
„ echinobothrida in Indian
poultry . 163 I
„ tetragona in Indian poultry 163
Davey, J. B., and Newstead, R. Mos¬
quitoes and other blood-sucking j
arthropods of the Upper Shiri
River, Nyasaland. 457
Diarrhoea, Monilia spp. recovered
from cases of.272, 275
Dicranotaenia spbenoides in Indian
poultry. 163
Dicrobezzia nigritibialis , sp.n., adult 371 1
» „ » pupa 373 !
Dilepis kempi , sp.n., from a cormorant 169
Diorchis jlavescens in a new host . 403
„ sp. in Indian poultry . 165
Dragonflies destructive to S. calopus... 307 ,
Dysenteric diarrhoea, Monilia sp.
recovered from a case of. 272
E chesty pus sepiaceus . 461
Effect of saline solution and sea-water
on S.fasciata . 377
Entamoeba coli cysts, Sizes of, in
symptomless carriers 149
„ „ differentiation from E.
histolytica . 121
„ „ Incidence of, in symp¬
tomless carriers... 120, 123
„ „ irregular distribution in
a stool... 118
„ histolytica cysts, Sizes of, in
symptomless
carriers. 133
„ „ danger of spread
by healthy |
carriers. 126
„ „ differentiation
from E. coli ... 121
„ „ Incidence of, in
symptomless '
carriers ...120, 124
„ „ spread by flies... 126
Entomological research by the Liver¬
pool School . 38
Etorleptiomyia mediolineata . 460
Eukraiohelea , gen. nov. 347
PAGE
Eukraiohelea afrtcana , sp.n., adult ... 347
„ versicolor , sp.n., adult ... 351
Euproctus-biceronatum group of Cyli-
costomum . 400
Evans, A. M. Notes on Culicidae
collected in Venezuela . 445
Evans, A. M., and Newstead, R. New
tsetse-flies ( Glossina ) from the Bel¬
gian Congo . 95
Evans, A. M., and Newstead, R.
Report on rat-flea investigation.. .... 287
Feeding habits of S. calopus . 265
Fleas occurring on rats at Liverpool:.. 287
Flies as carriers of E. histolytica . 126
Filariasis research by the Liverpool
School . 37
Fungal infections in West Africa . 271
Fungus of the genus Nocardia culti¬
vated from heart blood. 283
Giardia intestinalis , even distribution
in a stool. 118
„ „ Incidence of, in
symptomless
carriers ...120,127
Glossina , new, from the Belgian Congo 95
99
fusca y genital armature .
102
99
„ var. congplensis y var.
nov.
99
99
„ var. congolensis , var.
nov., genital arma-
ture .
102
99
schwetzi , sp.n., genital arma¬
ture.
99
99
99 99 morphology
95
99
tabaniformis , genital armature
99
Gold Coast, Bronchomoniliasis in...53, 285
,, ,, Ceratopogoninae of.... . l 77
Gordon, R,. M., and Young, C. J.
The feeding habits of Stegpmyia
calopus . 265
Granuloma, ulcerative, Notes on. 413
Gyrocoelia genus, Diagnosis of . 412 4
Fielminthiasis research by the Liver¬
pool School...,. 36
Hemiptera from Nyasaland. 461
Heteroplus grandis from an Indian
jungle fowl . 166
Hill, G. F. Musca dome stir a as a
4 bush-fly * in Australia . 93
Hill, G. F. Notes on some unusual
breeding-places of Stegpmyia fas data
in Australia. 91
vi
PAGE
Hongkong, Sanitary and housing con¬
ditions in . 219
„ Tuberculosis in ...213, 227, 381
Hookworm-infected children, Arneth
count in . 49
Horses, Lappeted Anoplocephala in ... 249
Hyalomma aegyptium . 462
Ihle, J. E. W. On the genus Cyli-
costomum . 397
India, Pappataci flies from . 103
Indian poultry, Cestodes from . 161
Indigenous infection with P. falciparum 59
Ingram, A., and Macfie, J. W. S.
Bronchomoniliasis complicating pul¬
monary tuberculosis in a native of
the Gold Coast .53, 285
Ingram, A., and Macfie, J. W. S. A
fungus of the genus Nocardia culti¬
vated from heart blood.. 283
Ingram, A., and Macfie, J. W. S.
West African Ceratopogoninae. 313
Ingram, A., Carter, H. F., and Macfie,
J. W. S. Observations on the Cera-
topogonine midges of the Gold
Coast, with descriptions of new
species . 177
Jngramia uniformis . 460
Intestinal parasites among symptom¬
less carriers in Jamaica . 117
Isle of Man, Mosquitoes in . 73
Ixodidae from Nyasaland . 461
Jamaica, Sizes of E . colt cysts in symp¬
tomless carriers in 149
„ „ E. histolytica cysts in
symptomless car¬
riers in . 133
„ Symptomless carriers of in¬
testinal parasites in. 117
yoblotia digitatus . 446
Kempia ochrosoma , sp.n., adult . 340
„ larva . 343
„ pupa . 34 2
Kennan, R. H. Multiple aneurysms
in a child . 245
Knapp’s observations on Arneth count
in hookworm patients . 49
Krusei group of Monilia . 279
Lappeted Anoplocephala in horses ... 249
Leishmaniasis treated with tartar
emetic solution. 107
Leprosy, Case of suspected. 173
„ Note on a case of. 269
PAGE
LeptopsyUa musculi on rats at Liverpool 293
Ltbellulidae destructive to S. calopus... 302
Liverpool, Fleas occurring on rats at... 287
„ School of Tropical Medi¬
cine : Scientific Record 1
Macfie, J. W. S. The effect of saline
solutions and sea-water on Stegomyia
fasciata . 377
Macfie, J. W. S. Notes on 6ome
fungal infections in West Africa. 271
Macfie, J. W. S., and Ingram, A.
Bronchomoniliasis complicating pul¬
monary tuberculosis in a native of
the Gold Coast .53, 285
Macfie, J. W. S., and Ingram, A. A
fungus of the genus Nocardia culti¬
vated from heart blood. 283
Macfie, J. W. S., and Ingram, A.
West African Ceratopogoninae. 313
Macfie, J. W. S., Carter, H. F., and
Ingram, A. Observations on the
Ceratopogonine midges of the Gold
Coast, with descriptions of new
species . 177
Macfie’s observations on Arneth count
in hookworm patients. 50
MacGregor, M. E. The structural
differences in the ova of Anopheles
maculipennis, A. bifurcatus , and A.
plumb eus . 417
l Malaria, case of indigenous malignant
| tertian . 59
„ on a Venezuelan oilfield. 435
| „ research by the Liverpool
; School . 5
Malignant tertian malaria, case of
indigenous...... 59
Mansonioides unijormis . 459
Maplestone, P. A. Notes on Austra¬
lian Cestodes.403, 407
Maplestone, P. A. Notes on ulcerative
granuloma . 413
Maplestone, P. A., and Southwell, T.
A note on the synonymy of the
genus ZschokkeeUa , Ransom, 1909,
and of the species Z. guineensis ,
| (Graham, 1908) . 455
i Metroliasthes lucida in Indian poultry 163
Midges of the Gold Coast. 177
Moniezia alba in a new host . 405
Monilia , Castellani and . Chalmers’
classification of.56, 273, 278
PAGE
Monilia , species of the Krusei group... 279
„ „ „ Pseudolondinen -
sis group ... 277
„ „ „ Tropicalis
group......57, 274
„ accrarnstSy sp.n., Biochemical
reaction
of. 57 . 274
„ „ „ from a case
of tuber¬
culosis, 55, 274,
28 5
„ afncana , sp.n., Biochemical
reaction of 277
„ „ „ from a case of
diarrhoea... 275
„ enteric a y Biochemical reaction
of .;.;- 57 . 274
„ enterocola , sp.n., Biochemical
reaction of 279
„ „ „ from a case of
diarrhoea.. 277
„ insolita. Biochemical reaction
of.•••:.;.:- 57 .274
„ krusei , Biochemical reaction of 279
„ me ta tropic alts. Biochemical
reaction of .57, 274
„ nrp/d, Biochemical reaction
of .:.;.;. 57 . 274
„ parakrusei 9 Biochemical re¬
action of. 279
,, paratropicalis. Biochemical
reaction of .57, 274
„ pseudolondinensis , Biochemical
reaction of. 277
„ pseudolondinoides. Biochemical
reaction of .. 277
„ pulmonalis , Biochemical re¬
action of .57, 274
„ tropicalisy Biochemical re¬
action of .57, 274
Monohelea litoraureay sp.n., female. 344
Monopylidium gaUinarum y sp.n., in *
Indian poultry. 164
Montgomeryi group of Cylicostomum ... 401
Mosquitoes, Apparatus for the in¬
dividual breeding of ... 473
„ in the Isle of Man. 73
♦, and other blood-sucking
arthropods of the
Upper Shiri River,
Nyasaland . 457
PACK
Mosquito ova, Certain uniform charac¬
ters of. 418
| Multiple aneurysms in a child . 245
Musca domestica as a ‘ bush-fly * in
Australia . 93
Newstead, R., and Davey, J. B.
Mosquitoes and other blood-sucking
arthropods of the Upper Shiri River,
Nyasaland . 457
Newstead, R., and Evans, A. M. New
j tsetse-flies (< Glossina ) from the Bel-
| gian Congo . 95
Newstead, R., and Evans, A. M.
Report on rat-flea investigation. 287
Newstead, R., and Sinton, J. A. On
a collection of Pappatad flies
( Phlebotomus ) from India . 103
Nocardia cruoris , sp.n., cultivated from
| heart blood . 285
I Nocardiomycosis, tonsillar, in West
I Africa . 282
Nyasaland, blood-sucking arthropods
of the Upper Shiri River . 457
Olfersia ardeae . 461
Ophiotaenia bylae in a new host . 404
„ longmani . 404
Oriental sore treated with tartar
emetic solution. 107
Otomycosis, Note on a case of. 279
Pantala fiavescens destructive to 5 .
calopus, . 303
„ „ Feeding experiments
| with . 304
Pappatad flies from India . 103
Parasites, intestinal, among symptom¬
less carriers in Jamaica . 117
Phenvlglycine amido arsenate of
sodium, Effect of, on T. brucei in
rats and T . rhodesiense in mice . 427
1 Phlebotomus antennatus from India ... 105
„ major from India . 105
„ minutus from India . 104
var. antennatus
from India 105
„ africanus from
Nyasaland 460
„ papatasii from India . 104
„ sergenti from India . 105
Piroplasmosis research by the Liverpool
School . 16
Pistia stratiotesy source of mosquitoes
and midges . 314
viii
Plajmodium falciparum , Indigenous
infection
PAGE
Scientific Record of the Liverpool
School of Tropical Medicine .
PAGE
with. 59 1
Amoebiasis..
34
99
„ infection, ;
Beri-beri .
35
incubation
Blackwater Fever.
H
period, 62,63,68 !
Entomology.
38
99
„ infection, (
I Filariasis.
37
incubation
Helminthiasis .*.
36
period
Malaria and Sanitation .
5
following
Piroplasmosis ..
16
inoculation, 64,65
Protozoology ...
45
99
„ " infection,
Relapsing Fever and Spiro-
incubation
chaetes .
3 i
period
Trypanosomiasis .
16
following
Yellow Fever.
28
bite .66, 67
1 Scott, H. H. The incidence
of
Poultry, Indian, Cestodes in. 161
Prevalence and character of tubercu¬
losis in Hongkong.213, 227, 381
Probezzia pistiae , sp.n., adult . 364
n » » larva. 368
„ » » pupa . 367
„ stepbensi, sp.n., male . 369
Protozoa, Symptomless carriers of. 117
Protozoological research by the Liver¬
pool School. 45
Pseudolondinensis group of Monilia ,
Biochemical reactions of. 277
Psotophora posticata . 446
Psychodidae from Nyasaland. 460
Pupipara from Nyasaland . 461
Queensland, Arneth count in hook-
worm-infected children in. 49
Radiatum-elongatum group of Cylicos -
tomum . 399
Rat-flea investigation, Report on. 287
Rats, African, Cestodes from . 167
Relapsing Fever research by the Liver¬
pool School. 31
Rhipecephalusfalcatus . 462
„ maculatus . 462
„ neavei . 461
„ simus . 462
Rhodesian sleeping sickness treated
with * Bayer 205 * . 479
Russian Turkestan, Oriental sore in ... 107
Saline solutions, Effect of, on S.fas data 377
Scbizodactylus, gen. nov. 353
„ Ulmatoscopus, sp.n.,
adult 353
,, ,, sp.n.,
pupa... 356
149
>33
intestinal parasites, especially with
regard to the protozoa, amongst
symptomless carriers in Jamaica. 117
Scott, H. H. The prevalence and
character of tuberculosis in Hong¬
kong .213, 227, 381
Scott, H. H. A study of the sizes of
Entamoeba colt cysts amongst
symptomless carriers in Jamaica
Scott, H. H. A study of the sizes of
Entamoeba histolytica cysts amongst
symptomless carriers in Jamaica
Sea-water, Effect of, on S.fasciata ... 377
Sinton, J. A. Notes on Oriental sore
in Russian Turkestan and the results
of treatment with injections of
tartar emetic solution. 107
Sinton, J. A., and Newstead, R. On
a collection of Pappataci flies
(Phlebotomus ) from India . 103
Sleeping sickness, case of Rhodesian,
treated with ‘ Bayer 205 * . 479
Southwell, T. Cestodes from Indian
poultry. 161
Southwell, T. Cestodes from African
rats. 167
' Southwell, T. A new species of
Cestoda from a cormorant . 169
Southwell, T., and Maplestone, P. A.
A note on the synonymy of the genus
Zschokkeella y Ransom, 1909, and of
the species Z. gutneensis , (Graham,
>908).
Southwell, T., and Yorke, W. Lap-
peted Anoplocephala in horses . 249
Sphaeromias litoraurea } sp.n., male ... 359
455
IX
PAGE
PAGE
Spirochaetes, Research on, by the
Liverpool School. 31
Spleen in indigenous P. falciparum
infection.60, 63
Sprue-like diarrhoea, Cases of, at Accra 275
Stegomyia calopus y Feeding habits of.... 265
„ „ Natural enemies of 301
„ fasciata , Effect of saline solu¬
tions and sea¬
water on. 377
„ „ found breeding in
rot-holes in
Australia . 91
Stephens, J. W. W. Malaria on a
Venezuelan oilfield . 435
Stephens, J. W. W., and Adler, S.
A case of suspected leprosy. 173
Sterigmatocystis antacustica from cases
of otomycosis. 280
„ nidulans from cases of
otomycosis . 280
„ sp. in a case of otomy¬
cosis at Accra. 280
Symptomlcss carriers of intestinal para¬
sites . 117
„ „ Sizes of E. coli
cysts in. 149
„ „ Sizes of E. histo¬
lytica cysts in 133
Tabanidac from Nyasaland . 460
Tahanus africanus . 460
„ taeniola var. variatus . 460
Taenia magna . 255
„ plicata . 255
„ zehrae , Collin . 256
„ „ Sander. 255
Taeniorhynchus titillans . 445
Tartar emetic in treatment of Oriental
sore . 107
Tetracantbum-coronatum group of Cyli-
costomum . 397
Theobaldia annulata in the Isle of
Man.75, 88
Thysanosoma giardi in a new host. 405
Tonsillar nocardiomycosis, Case of, in
West Africa . 282
Trichiuris in symptomless carriers ... 123
Trichonema genus, question of nomen¬
clature (footnote). 397
Trigonotylus brevipes . 461
Tropicalis group of Monilia , Bio¬
chemical reactions of .57, 274
Trypanocidal effect of bismuth . 433
„ effect of phenylglycine
amido arsenate of
sodium on T. brucei in
rats and T. rhodesiense
in mice. 427
Trypanosome brucei in rat9 treated with
phenylglycine amido
arsenate of sodium ... 427
„ rhodesiense , Case infected
with, treated with
‘ Bayer 205 9 . 479
„ rhodesiense in mice treated
treated with phenyl¬
glycine amido arsenate
of sodium . 427
Trypanosomiasis research by the Liver¬
pool School. 16
Tsetse-flies, new, from the Belgian
Congo . .. 95
Tuberculosis complicated by broncho-
moniliasis in the Gold
Coast.53, 285
„ Education as a weapon
against. 224
„ in Hongkong ...213, 227, 381
„ Morbid anatomy in cases
of . 381
„ Portals of entry and mode
of spread of, 214, 219, 227
„ Possible congenital infec¬
tion with . 215
„ Racial susceptibility to... 218
Ulcerative granuloma, Notes on . 413
Unusual breeding-places of Stegomyia
fasciata in Australia. 91
Venezuela, Notes on Culicidae col¬
lected in... 445
Venezuelan oilfield, Malaria on a. 435
Water-bugs destructive to S. calopus
larvae . 303
West Africa, Fungal infections in....... 271
West African Ceratopogoninae ...177, 313
Xenopsylla cheopis on rats at Liverpool 290
„ „ breeding-places of... 295
Yellow Fever research by the Liverpool
School . 28
Yorke, W. Treatment of a case of
Rhodesian sleeping sickness by the
preparation known as ‘ Bayer 205 ’ 479
Yorke, W., and Adler, S. Note on a
case of leprosy. 269
x
PAGE
Yorke, W., and Southwell, T. Lap-
peted Anoplocephala in horses. 249
Young, C. J. Natural enemies of
Stegomyia calopus . 301
Young, C. J., and Gordon, R. M.
The feeding habits of Stegomyia
calopus . 265
PAGE
Zaitha spp. destructive to 5 . calopus
larvae . 303
Zebra, Anoplocephala rhodesiensis in.... 249
Z*schokkeella genus, Synonymy of. 455
„ guineensis from an African
rat . 167
„ „ Synonymy of.... 455
INDEX OF GENERA, SPECIES AND VARIETIES NEW TO SCIENCE
PAGE !
Angularia australis . 407
Atrichopogon africanum . 334
Atricbopogon chrysosphaerotum . 337
Atrichopogon elektrophaeum .;... 335
Atrichopogon homoium . 338
Atrichopogon perfuscum . 337
Bez&ia foyi . 361
Culicoides corsoni . 324
Culicoides inomatipennis , var. rutilus. .. 326
Culicoidcs nigeriae . 325
Cylicostomum labia turn, var. digitatum 401
Dasyhelea bootbi . 331
Dasybelea Jlava . 196
Dasyhelea flaviformis . 201
Dasyhelea fusca . 204
Dasyhelea fusciformis .. 209
Dasyhelea fuscipleuris . 200
Dasyhelea fusciscutellata . 187
Dasyhelea inconsptcuosa . 191
Dasyhelea luteoscutellata . 191
Dasyhelea nigeriae . 329
Dasyhelea nigricans . 194
Dasyhelea nigrofusca .
Dasyhelea paUidihalter .
Dasyhelea retorta .
Dasyhelea similis .
Dicrobezzia nigritibialis .
Dilepis kempi .
Eukraiohelea genus .
Eukraiohelea versicolor .
Glossina fusca, .var. congolensis.
Glossina schwetzi .
Kempia ochrosoma .
Monilia accraensis .
Monilia africana .
Monilia enterocola .
Monohelea litoraurea .
Monopylidium gallinarum
Nocardia cruoris .
Probezzta pistiae .
Probezzta stephensi .
Schizodactylus genus .
Scbizodactylus telmatoscopus .
Spbaeromias litoraurea .
k
PAGE
20 7
I84
333
189
371
169
347
35 1
99
95
34 °
285
277
278
344
164
285
364
369
3 S 3
353
359
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