The Frfsh-Vater Ijollusca of the Tanganyika Territory
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THE FRFSH-VATER IJOLLUSCA OF THE TANGANYIKA TERRITORY AND ZANZIBAR PROTECTORATE, AND OTHER CONTRIBUTIONS TO THE SYSTEATICS BIONO1ICS AND DISTRIBUTION OF FRESH-WATER AND TERRESTRIAL L1OLLUSCA. THESIS Submitted for the DEGREE of D.Sc. in The University of Edinburgh by WALTER ALAN MOZLEY, Ph.D.(Edin.) April 1940. CONTENTS List of Publications by Walter Alan tIozley. Papers: 1. The Freshwater Mollusca of the Tanganyika Territory and Zanzibar Protectorate, and their Relation to Human Schistosomlasis. Trans. Roy. Soc. Edinburgh, LIX,(No.26), 1939- A Biological Study of the Sub-Arctic Uollusca. Proc. Amer. Philos. Soc., 78, 1, 1937- The Fresh-Water Mollusca of Sub-Arctic Canada. Canadian Jour. Research, D,16 2 1938- 1 . PUBLICATIONS by WALTER ALAN MOZLEY 1925 Segmeritina crassilabris Walker in Manitoba. Canadian Field Naturalist, 39, p.85 1926 Molluscs from the Manitoba-Ontario Boundary. Nautilus, 39, pp.121-128. Preliminary List of the Mollusca of Jasper Park, Alberta. Nautilus, 40, pp.53-56. Some Molluscs from Western Canada. Nautilus, 40 0 pp.56-63. Molluscs from the Lake Brereton District, Manitoba. Canadian Field Naturalist, 41 0 pp.59-61. 1927 Some Plancton Organisms from Lake Brereton, Manitoba. Canadian Field Naturalist, 41, pp.84-85. 1928 The Variation of Lymnaea traskii Tryon In Pond and Lake Habitats. American Naturalist, 42, pp.286-288. Post Glacial Fossil Mollusca from a Delta Deposit near Winnipeg, Manitoba. Geological Magazine, 65, pp.267-270. New Records of Western Canadian Mollusca. Nautilus, 42, pp.13-18. 104 Note on Some Fresh Water Mollusca inhabiting Temporary Ponds in Western Canada. Nautilus, 42, pp.19-20. 1930 Further Records of Western Canadian Mollusca. Nautilus, 43, pp.79-85. Reports of the Jasper Park Lakes Investigations 1925-26. The Mollusca of Jasper Park. Transactions Royal Society Edinburgh, 56, pp. 647-669. 1931 Fresh Water Molluscs in Some Jasper Park Lakes. Canadian Alpine Journal, 1931, pp.148-158. 1932 A Biological Study of a Temporary Pond in Western Canada. American Naturalist,, 66, pp.235- 249. A New Interglacial Pulmonate Mollusc from the Province of Saskatchewan. American Midland Naturalist, 13, pp.236-240. 2 . Notes on Western Canadian Mollusca - Planorbis campanulatus wisconsinensis Winslow. Nautilus, 46, pp.59-62. 1933 The Local and Geographical Distribution of Some Rocky Mountain Mollusca. Proceedings Malacological Society London, 20, pp.214-221. A New Variety of Lymnaea from the Rocky Mountain Region of Canada. Proceedings Malacological Society London, 20, pp.241-243. 1934 The Discovery of Aearithinula Say in Central Siberia. Nature, 133, Molluscs from the Provinces of Saskatchewan and Alberta, Canada. Proceedings L'Lalacological Society London, 21, pp.138-145. x New Fresh-water Molluscs from Northern Asia. Smithsonian Misc. Collections, 92, No.2. Post Glacial Fossil Molluscs from Western Canada. Geological Magazine, 71, pp.370-382. x 1935 The Fresh-water and Terrestrial Moilusca of Northern Asia. Trans. Royal Society, Edinburgh, 58, pp.605-695. The Variation of two Species of Lymnaea. Genecics, 20, pp.452-465. 1936 The Statistical Analysis of the Distribution of Pond Molluscs in Western Canada. American Naturalist, 70, pp.237-244. 1937 The Ponds, Lakes and Streams of the Kirghiz Steppe. Scottish Geographical Magazine, 53, PP-1-10. The Status of the Greenland Lymnaeae. Proceedings Malacological Society London, 22, pp.186-188. The Terrestrial Mollusca of Sub-Arctic Canada. Proceedings Malacological Society London, 22 2 pp.38-382. A Biological Study of the Sub-Arctic Mollusca. Proc. American Phil. Soc. 78, pp.147-189. Frozen Ground in the Sub-Arctic Region and its Biological Significance. Scottish Geographical Magazine, 53, pp.266-270. 3l./ 3. 1938 The Fresh-Water i1o11usca of Sub-Arctic Canada. Canadian Jour. Research, 16, pp.93-138. The Sub-Arctic Region as a Molluscan Habitat. 1ature, 142, pp.1116-1117. 1939 The Quill Lakes Basin, Saskatchewan, Canada, and its Molluscan Fauna. Internat. Rev. Hydrol. u. Hydrobiol., 38, pp.243-249. The Fresh-water Mollusca of the Tanganyika Territory and Zanzibar Protectorate, and their Relation to Human Schistosomjasjs. Trans. Royal Society Edinburgh, 59, pp. 687-744. The Variation of Lyinnaea stagnalis (Linné). Proceedings Lialacological Society London, 23, itg-q, X Note: Nos. 21 and 23 only were submitted as Thesis for the degree of Ph.D. - TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH. VOL. LIX—PART III—(No. 26). 1938-1939. IHE FRESH-WATER MOLLUSCA OF THE TANGANYIKA TERRITORY AND ZANZIBAR PROTECTORATE, AND THEIR , RELATION TO HUMAN SCHISTOSOMIASIS. BY ALAN MOZLEY, PH.D.(EDIN.), WANDSWORTH SCHOLAR OF THE LONDON SCHOOL OF HYGIENE AND TROPICAId MEDICINE. [WITH THREE PLATES AND TEN TEXT-FIGURES.] EDINBURGH: PUBLISHED BY ROBERT GRANT & SON, LTD., 126 PRINCES STREET, AND WILLIAMS AND NORGATE, LTD., 36 GREAT RUSSELL STREET, LONDON, W.C. 1. AND PRINTED BY NEILL & CO., LTD., 212 CAUSEWAYSIDE, EDINBURGH, 9. MCMXXXIX. Price Eight Shillings and Sixpence. ( 687 ) XXVI.—The Fresh-water Mollusca of the Tanganyika Territory and Zanzibar Protectorate, and their Relation to Human Schistosomiasis. By Alan Mozley, Ph.D.(Edin.), Wandsworth Scholar of the London School of Hygiene and Tropical Medicine. (With Three Plates and Ten Text-figures.) (MS. received December 2, 1938. Revised MS. received March 29, 1939. Read March 6, 1939. Issued separately July 6, 1939.) CONTENTS. PAGE PAGE I. INTRODUCTION . . . . . 687 V. MEANS 01? CONTROL . . . . . 717 II. CENTRAL AFRICAN MOLLUSCA IN RELATION TO Drainage; Filling-in; Flushing; Chemical HUMAN ScmsTosolalAslS . . . 687 Treatment; Removal of Vegetation; Pro- III. SYSTEMATIC ACCOUNT OF THE MOLLUSCAN FAUNA 689 motion of the Growth of Vegetation; Enemies IV. THE DISTRIBUTION OF FRESH-WATER MOLLUSCA of Mollusca; Education. IN TANGANYIRA AND ZANZIBAR . . 706 VI. REPRESENTATIVE MOLLUSCAN HABITATS . . 727 i. Geographical Conditions which Affect VII. SUMMARY AND CONCLUSIONS . . . 741 Molluscan Distribution . . . 706 ii. Local Conditions which Affect Molluscan VIII. REFERENCES TO LITERATURE . . 743 . . Distribution 707 ix. DESCRIPTION OF PLATES . . 743 iii. Types of Aquatic Environment . 714 I. INTRODUCTION. During a part of the years 1937 and 1938 a biological study of the fresh-water mollusca of the Tanganyika Territory and Zanzibar Protectorate was carried out with the aid of the grant of the Wandsworth Scholarship of the London School of Hygiene and Tropical Medicine. Up to the present little has been known about the molluscs of this part of Africa, although the importance of certain members of this group as the intermediate hosts of human blood flukes (Schistosoma spp.) has been widely recognized. The following account of the constitution and distribution of the molluscan fauna of this region may therefore be of interest and value. The author is indebted to Dr R. R. Scott and to Dr W. L. Webb, Directors of Medical Services in Tanganyika and Zanzibar respectively, and to their subordinates for their active co-operation in the course of this investigation. The interest, taken in the work by Mr C. E. Bobbins, General Manager of the Tanganyika Railways, and by Dr C. Gilman, formerly Chief Engineer of those railways, and now Water Consultant to the Tanganyika Government, has also been of great aid. Major M. Connolly has given valuable assistance in the preparation of the systematic section of this paper. The author is also greatly indebted to Professor B. T. Leiper, F.R.S., for his constant interest in this work, and to Mr W. A. McDonald for valuable suggestions and aid in many ways. A grant of £50 towards the cost of publication has been given by The London School of Hygiene and Tropical Medicine. H. CENTRAL AFRICAN MOLLUSCA IN RELATION TO HUMAN SCHIsTosoMIAsIs. At least two species of blood flukes parasitic in man are known to occur commonly in Central Africa. Schistosoma hcematobium is found in Zanzibar, Pemba, Tanganyika, and the adjoining territories. Schistosoma mansoni is widely distributed on the African mainland but is not native to Zanzibar and Pemba Islands. Roughly speaking, from 10 to 70 per cent. of the native population of infected districts harbour these parasites. The Director of Medical TRANS. ROY, SOC, EDIN., VOL. LIX, PART III, 1938-3 (NO. 26). 104 688 DR ALAN MOZLEY ON FRESH-WATER MOLLTJSOA OF TANGANYIKA TERRITORY Services of Zanzibar in his Annual Report for the year 1935 (p. 16) made the following state- ment regarding schistosomiasis, " The disease treatment figures for this disease, 692 males and 147 females, give no idea of the extreme prevalence of it among the children of Pemba. In many of the villages in low-lying valleys, or in the neighbourhood of streams and swamps in Pemba, passage of a little bloody urine towards the end of micturition is universal among the boys, and those boys without obvious blood in the urine generally show albumen or even eggs on investigation." Further, in the same report (p. 39), the condition in certain localities is described more precisely. "It will be observed that the percentage of pupils infested with this parasite (S. hcematobium) is extremely high (81-25 per cent. in Nwachani and 7749 per cent. in Kengeja schools). The percentages given in these tables for this helminth must, as in the case of the intestinal parasites, be taken as the absolute minimum, because re-examina- tion of the urines of these children which are found to be negative on the first examination may prove positive on subsequent examination." For the most part human schistosome infections in this part of Africa are light as com- pared with those found in Egypt. The medical importance of these light infections has been emphasized by Blackie (1932). At the present time it is not possible to evaluate the economic importance of human schistosomiasis in this sector of Africa, but it is doubtless very considerable. The local intermediate host of Schistosoma mansoni is the snail, Biomphalaria pfeifferi Krs. Immature specimens of this schistosome were obtained from the liver of a monkey (Gercopithecus sp.) which had been experimentally infected with cercaria obtained from "wild" Biomphalaria pfeifferi collected near Mwanza, T.T.