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This electronic thesis or dissertation has been downloaded from Explore Bristol Research, http://research-information.bristol.ac.uk Author: Sen, B Title: Phytoplankton of Shearwater, with special consideration of fungal parasites and epiphytes General rights Access to the thesis is subject to the Creative Commons Attribution - NonCommercial-No Derivatives 4.0 International Public License. A copy of this may be found at https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode This license sets out your rights and the restrictions that apply to your access to the thesis so it is important you read this before proceeding. Take down policy Some pages of this thesis may have been removed for copyright restrictions prior to having it been deposited in Explore Bristol Research. However, if you have discovered material within the thesis that you consider to be unlawful e.g. breaches of copyright (either yours or that of a third party) or any other law, including but not limited to those relating to patent, trademark, confidentiality, data protection, obscenity, defamation, libel, then please contact [email protected] and include the following information in your message: •Your contact details •Bibliographic details for the item, including a URL •An outline nature of the complaint Your claim will be investigated and, where appropriate, the item in question will be removed from public view as soon as possible. PHYTOPLANKTON OF SHEARWATER WITH SPECIAL CONSIDERATION OF FUNGAL PARASITES AND EPIPHYTES A thesis submitted for the Degree of Doctor of Philosophy at the University of Bristol by Bülent Sen June 1982 MEMORANDUM The work reported ýn this thesis is the result of my own independent research under the super- vision of Professor F. E. Round, University of Bristol. /J ,. ---r ý.ý_v . ý*, ý i. ACKNOWLEDGMENTS I am very grateful to Professor F. E. Round for supervising the research, for helping me with English and for his advice and encouragement throughout this work. I would like to thank Dr. R. M. Crawford for his technical advice and constructive discussions, and Dr. M. F. Madelin for his discussion on the fungal parasites. I am also indebted to the staff of the department for their, help and kindness. I would also like to thank Mr. Tim Colborn for preparing the illustrations and_Mrs. Gillian Lockett for typing the manuscript. My thanks are also due to Mr. Sabri Erdem, Mr. Necdect Bayram, Mr. Oguz Coser and Mr. Tevgik Erkal for their photographic assistance. I also wish to acknowledge the moral support and encouragement given by the Misses Angela, Rachael and Suzi Kiely and my parents. Finally, I am grateful to the Turkish Government for providing the finance and for the opportunity to study in England. SYNOPSIS The phytoplankton of Shearwater, Wiltshire has been studied over three years paying particular attention to the epiphytic populations on the individual algal species. In order to relate any effects of the epiphytes on the seasonal growth of the algae it was necessary to monitor the physico- chemical factors and compare the cycles of individual algae and groups of algae with these factors. Unlike many more oligotrophic lakes, Shearwater maintained high populations of phytoplanktonic algae throughout the whole study period. Individual species also maintained their growth over longer periods than expected. Correlations between phytoplankton and physico-chemical factors were made wherever possible. Epiphytic populations of fungi (mainly Chytrids) protozoa and other algae were continually monitored on the phytoplankton. in the case of the Chytrids, this also involved studies of stages in their life cycles in order to identify the fungi. Their effects on the growth patterns of the algae were documented. There is no literature on the occurrence of choanoflagellates on algae and hence the present study of these in Shearwater provides unique data. Data on the epiphytic populations was obtained by a combination of light and electron microscopy. The taxonomic position of the epiphytes and their relationship to the phytoplankton is discussed with reference to the literature. One new fungal species Is described. CONTENTS Introduction 1 Chapter 1. Introduction to Shearwater, methods and physico-chemical data. 3 , 1. Introduction 3 2. Ecological studies 4 3. Methods and field work 4 4. Physico-chemical features 9 Chapter 2. Phytoplankton 22 1. Seasonal cycles of BacLllariophyceae 24 2. Seasonal cycles of Cyanophyceae 59 3. Seasonal cycles of Chlorophyceae 64 Chapter 3. Fungi 82 1. Review of previous work 84 2. Methods 104 3. Parasitism of Bacillariophyceae 105 4. Parasitism of Chlorophyceae 162 5. Parasitism of Cyanophyceae 182 Chapter 4. Colourless Epiphytes 193 Chapter S. Concluding discussion 214 References 219 1. INTRODUCTION A considerable amount of information has been gathered since the beginning of the present century in limnological studies of many waters of the world. The bulk of these studies have invariably concentrated on the seasonal periodicity of phytoplankton, primary productivity and water chemistry. Physico-chemical factors have generally been considered primarily responsible for the seasonal cycle of planktonic algae, consideration of biological factors, such as parasitism and interaction between organisms being largely ignored. In fact, very few investigations have been made hitherto on the effect of parasitic fungi on phytoplankton populations despite the fact that pioneer works of CANTER & LUND (1948,1951) showed that these organisms could exert an important influence on the cycle. Algae form the major part of phytoplankton and include both eucaryotic and procaryotic organisms which may be parasitized by viruses, bacteria, fungi and protozoans. In fact, organisms in aquatic environments and indeed in other communities are interdependent in many ways and their distribution should not be interpreted without considering their mutual inhibitory and stimulatory effects. The present work comprises a comprehensive study of the effects of fungal parasitism together with physico-chemical factors on the seasonal cycle of phytoplankton population in a freshwater lake. It is also well known that some algae, bacteria and protozoans often occur on algal cells or in the mucilage surrounding them. It is yet not known to what extent these non-parasitic micro- Z. organisms affect the growth of algae. Shearwater -a freshwater lake investigated in this study - has provided good examples of this kind of micro-association during this study. The ecology of epiplanktonic organisms such as Stylosphaeridium stipitatum and members of the zoomastigo- phoran genera Bicosoeca, Codosiga and Salpingoeca has been studied quantitatively for the first time and the role of planktonic algae in harbouring significant populations of these organisms has been considered. In order to assess these relationships, it was essential to fully understand the seasonal periodicity of the representatives of the phytoplankton. 3. CHAPTER I. INTRODUCTION TO SHEARWATER - METHODS & PHYSICO-CHEMICAL'DATA Shearwater is a small, artificial lake in Wiltshire (Map ref. ST 850421) lying on Cretaceous Upper Greensand. It lies in a catchment of deciduous woodland and agricultural land and is therefore richly supplied with nutrients from small streams. It is used for recreation, mainly sailing. No detailed survey of the lake has been undertaken and only three papers on its algal flora have been published. ROUND (1965) studied the epipsammic flora of the sandy sediments, MOSS & ROUND (1967) reported on the standing crops of the epipelon and epipsammon and HICKMAN & ROUND (1970) studied the primary production of the epipelon and epipsammon over a two year period. The rich phytoplankton has never been sampled before and so this is a first account both of the phytoplankton, its associated epiphytes and parasites and the chemical status of the water. Sampling was from a wooden platform built out into the water for use of the sailing club and hence all the data has been obtained on sub-surface populations close to the shore. No information is available on changes with depth or of the sequence of stratification of the water. The flora clearly indicates a eutrophic lake with a distinct seasonal cycle of spring diatoms, summer Chlorophyta and autumn Cyanophyta. The continual drainage and outflow over a weir maintains a relatively high nutrient supply throughout the year and the phytoplankton populations also tended to be high throughout the sampling period. 4. ECOLOGICAL STUDIES The literature on the ecology of freshwater phyto- plankton has now become columinous with the continuous studies of algal periodicity and the factors determining the ecology of phytoplankton. Many factors may influence the development of an algal bloom the duration of a bloom and the ultimate decline of a bloom in freshwaters. It is, therefore, difficult to review the effects of each environmental factor separately. The subject is so diverse that it cannot be dealt with on a simple fact to fact basis, but requires information from many fields of interest in order to understand this complex system. Some physical factors are very complex but it is usually easier to investigate the effects of physical factors than those of chemical factors. Biological and chemical factors merge into one another in the field of organic chemistry (e. g. extra-cellular and inhibitory substances). Nevertheless, field studies and data on the chemical composition of the aquatic environment are indispensible for the