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By Imperial Colleges London.. S, W, 7, October., 1954 ýl I STUDIES ON THE LIFE-HISTORY AND ECOLOGY . OF TREIMTODES OF THE GENUS RENICOLA COHN., 1904- Thesis submitted for the Degree Of Doctor of Philosophy in the University of London by Christopher Amyas Wrightv B. Sc, Imperial Colleges London.. S, W, 7, October., 1954. ST COPY AVAILA L Variable print quality Abstract Trenatodes of the genus Renicola from British, birds are described and evidence to show the relationship between these flukes and the "Rhodonotopa" group of cercariae is prosented. The fooding-habits of all of the SPOcies of birds recorded as hosts for Renicola in Western Europe. aro analyzed to give an indication of the probable second intermediate host in the cycle. The ecology of Turritella comunis is discussed and reasons for the relation between infection with sporocysts of the "Rhodometopa" Broup and the size of the snails are suggested. Attempts to infect Turritella connunis with eggs from Renicola sp. are described and the results discussed. The development of the sporocystqof the "Rhodometopat' group of corcariae within the interlobular spaces of the gonad of Turritella communis is described. A novi interpretation of the structure of the sporocyst body- wall is presented. The appearance of a second daughter generation of sporocysts is described. The Sporocysts Historical An examination of some of the literaturo on this stage in the development of trematodos reveals a con- siderable variance of opinion on the structure of sporocysts and the function of various organs and cells found in them. This is possibly due to the fact that most general works refer only to the mother sporoeyst and go into considerably more detail on the structure of rediae than of daughter sporocYsts. Baer (1951) states that sporocysts have no mouth or intestine and that some rediae which have a poorly developed pharynx and no intestine closely resemble sporocysts. on the other hand several sPOrocysts are described hs having 2.0rearia a pharynxp eog. s. pekinensis Faustp 1921 which has a "pharyngeal sphincter" but no gut, and the sporocysts of the "11hodometopa" group which, according to Rothschild (1935), have a pharynx "similar to that' of C. Faust, but surrounded by secretory cells _pekinensis the "salivary" resembling glands of rediao". I The histology of the sporocyst wall is an even more controversial subject. Stunkard (1932) states that the wall of the sporoo7st of Ccrcari_q rhodcm0topq consists Of two layers, an outer one packed with deeply (62) Contents Pago Introduction The Adult Flukes Historical 4 0.00 be be 04 Materials and, Methods Descriptions 14 Pathology, 27 32 Ecology and-Foeding of host species be The Mollucan Intermediate host 37 The Eggi., Mirqcidiun,, and, Infection experiments _, _48 The Sporoeysts Historical , 62 Materials and Methods 66 Development 67 .. - ý o. Mature'-daU'ghtor spprocysts 79 Effec, t,, on Turritella communis, 83 The Cercariae Historical 86 Ibe 04 SPecies found 04 00 00 Ob 88 be Cercaria doricha 00 89 Pipp-entata n. s; * Cercaria cooki n. sp, 92 Anatomy 06 so OF-7RE-odomatopall cercariae 96 Development .. 101 Shedding 04 00 015 *0 be 0000* 103 Behaviour 41 4.0 00 00 110 04 *6 of The Motacercariae Historical be 121 Species found 123 be be Origins and infection rates of Clupea sprattus 128 Infection 136 experiments be *0 00 0.6 Discussion 60 *0 'be 09 041 be 140 Appendix An attempt to raise some sea-birds 149 Sunmary . 00 00 *0 0* 0 00 00 154 References 00 as so be *0 be 157 Subsidiary Papers. 1. Probable relationship between the RhodometoDa group of cercariae and the Trematode genus Renicola Cobn. 2. Trematodes of the genus Renicola from the kidneys of birds in Brazil. 3. Trematodes of the genus Renicola from birds in British zoos with descriptions of two new species. Acknowledgments 11 .--ý, 1 I acknowledge with great thanks the supervision of Professor JJC. Buckley. 1,17 thanks are also due to Professor O. W. Richards in whose department this work has been carried out and to the Agricultural Research Council for the Scholarship which made this research possible. I an, unable to name all of those who have helped me in various ways, such as by the collection of material and by suggestions as to where and how "material might be obtained. The following are those to whom I am particularly indebted. Miss Mirian Rothschild., for the loan of material and for much helpful advice, 'including valuable suggest- ions as to possible errors in her own previous work. Mr. S. Prudhoe of the British Museum (Natural History)p for the loan of co-type material of-Renicola olandoloba'also for drawing my attention to some very recent literature and for his useful criticisms. Dr. T. G. Campbell, Poultry breeding unit of the British Enpire Cancer Campaigns Edinburghs for sections the of kidneys of Kine-peneuins infected with Renicola SP, Dr. Allen McIntosh of the Bureau of Animal Industries, Washington, for specimens of Renicola brantae. Dr. L. Harris on-Mathows R. R. S., Dr. 'W. C. Osman Hill and the staff of the-Prosectorium of the Zoological Society of-London for their interost and for much natorial. Dr. E. C. Appleby of the Royal Zoological SocietY of Scotland for supplying samples of droppings from the penguins in the Society's, menagerie at Edinburgh. -Mr. H. E. 'Jenndr'of, Lowe'stoft who for three winters has sent me oiled soa-birds washed up on the beachesp also Inspector P. Grinnallp R. S'. P. C. A. Bodnin.. Mr. W. E. Collins of Bognor Regis and Dr. Norman'lloore of'Bristol UniVerslty.. allýof whom have sent oiled birds to me. Mr. John Barrett and Mr. Peter Condor# wardens of Dale Fort Field Centro and Skokholn Island respectively for their help in many aspocts of tho fieldýwork and in particular for their assistance in obtaining Manx shearwaters. Mr. R. Smithers for his assistance in the attempt to raise some sea-birds in captivity. Mr. L. R. Brightwell of Peacehaven, Sussex for obtaining supplies of fish and arranging for trips with the Newhavon trawlers for me. Mr. A. C. Burd of the Ministry of Agriculture and Fisheries Laboratory, Lowestoft for samples of whitebait from, the East Coast.. also Mr. S. Young for smnples of Sprats from the Thames Estuary. Dr. N. Waloff and Dr. S. Markowski for their assistance in translating Russian literature. Finally, Vice-Admiral A. Day and Commanders G. S. Ritchie and G. P. D. Hall of the Hydroeraphic Dopartmenty Admiralty who made possible the dredging and plankton-netting work in St. Bride's Baty, also Dr. H. F. F. Herdnan of the National Institute of Oceanography who lent me equipment from R. R. S. Discovery II for this work and Commander R. H. Connell')-D. S. C., R. N. and the officers and men of H. Ivl*'S. Cook who'co-operated most readily in obtaining those samples. Introduction Describing the excretory system of Cercaria (193-2) rjga=Utora Perezp 1924, Stunkard wrote - "The system is so peculiar and characteristic that it will afford a quick and certain criterion for the identifica- tion of later stages in the life-history, and may,, indeed., lead to the correlation of this larva with an adult trematode having the same excretory pattern. " Such an adult trematode has now been found and the following is an account of some. observations on the ecology and development of both the cercariae and the adult flukes and of attempts to obtain expuimental evidence of the relations between them. In an earlier paper (Wright 1953) brief mention was made of the facts Which led to the undertaking of this work. k number of infections of Renicola in the kidneys of various birds in the London and other zoos were studied (Wright 1954P). of particular interest were the worms removed from three penguins. Campbell and Sloane (1943) also reported finding flukes of the genus Renicola in p enguins at the Edinburgh zoo. These authors stated that some of the penguins found to be infected had been hatched and raised in the menagerie. This fact and the finding of immature flukes together with very gravid spvcimens in the same bird in London led to the belief that the pprasite, was acquired from the food on which the birds were fed.. Enquiries showed that in London the principal food of the p.enguins is herring and whiting and in Edinburgh herring and haddock. This at once seemed to implicate herring as the likely vector and since most of the fresh herring sold in this country are caught fairly close to Britain it seemed probable that the paradite would occur in British sea-birds. A, further point in. favour of herring acting as the vector is that_the other two species of fishp both, Gadoidsg" are rather deep-water forms and not so readily caught by diving birds as the surface-feeding. 9 shoaling, Clupeoid, With the discovery of about ten Renicola sloanei in a Common guillemot from Pagham in Sussex the presence of the parasite in British birds was confirmed. Further work has since shown eleven further infections in three additional sp4es of hostso allZritish. Anatomical studies on the material from the penguins that the showed excretory vesicle in these flukes is of a peculiar, typev Y-shaped with numerous lateral branches and a cross-comection between the two arms of the Y (2) behind the ventral sucker. This seemed to be a suitable character by which any possibly related cercariae might be judged. Reference to the literature on cercariae from British marine gastropods soon showed that such a vesicle was to be found in the uRhodometopall group (Rothschild 1935).
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