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Universiteit O.V.S DJ.V.I. alBlIOTED UniVERSITEIT O.V.S. .. BIBllOTffK fERDIE EKSEMPLAAR MAG ONDER t University Free State }EEN OMSTANDlGHEDE UIT DIE 11111111111111111 ~IIIII~ IIIIIIII~ 1111111111111111111111111111111111111111111 34300001320336 3lRLlOTEEK VER\.VYDER WORD NIE Universiteit Vrystaat REPRODUCTIVE PROCESSES OF SCYPHIDIID PERITRICHS ASSOCIATED WITH LIMPET AND HALIOTID HOSTS ALONG THE COAST OF SOUTH AFRICA by Helene Peters Thesis submitted in fulfilment of the requirement for the degree Philosophiae Doctor in the Faculty of Natural and Agricultural Sciences, Department of Zoology and Entomology, University of the Free State Promotor Prof. Linda Basson Co-promotor Dr. Liesl L. van As November 2002 "Lord, you have made so many things! There isthe ocean, large and wide, Where countless creatures live, Large and small alike." Psalm 104:24 - 25 ITABLE OF CONTENTSI Chapter 1 INTRODUCTION 1 Chapter 2 MATERIALS AND METHODS 5 Study area 5 Collection of molluscs 5 Collection of symbionts 9 Preparation of material 9 Light microscopy 9 Hematoxylin 9 Mayer's Hematoxylin method 9 Harris' Hematoxylin method 10 Heidenhain's Hematoxylin method 11 Protargol 11 Scanning electron microscopy (SEM) 13 Morphological measurements 14 Authors of taxa 15 Terminology 16 Chapter 3 LITERATURE REVIEW: REPRODUCTION IN SOME PERITRICHS 25 ASEXUAL REPRODUCTION 25 Symmetrogenic Fission 25 Homothetogenic Fission 26 Multiple fission 26 Telotroch formation 27 Preconjugation fission 28 SEXUAL REPRODUCTION 29 Gametogamy 29 Autogamy 29,30 Gamontogamy 29 Anisogamonty 29 Conjugation 30 REPRODUCTIVE PROCESSES OF SOME REPRESENTATIVE CILIOPHORAN GENERA 31 BINARY FISSION 31 Binary fission in Rhabdosty/a vema/is Stokes, 1887 31 Binary fission in Urceo/aria synaptae Cuénot, 1891 32 Binary fission in Trichodina spheroidesi Padnos & Nigrelli, 1942 33 Binary fission in Scyphidia ameiuri Thompson, Kirkegaard & Jahn, 1947 34 Binary fission in Scyphidia micropteri Surber, 1940 34 Binary fission in Scyphidia tholiformis Surber, 1943 35 TELOTROCH FORMATION 35 Telotroch formation in Rhabdosty/a scyphidiformis Vávra, 1961 35 Telotroch formation in Circo/agenophrys ampulla (Stein, 1851) 36 Telotroch formation in Orbopercu/aria raabei Dobrzanska,1961 36 Telotroch formation in Scyphidia ameiuri Thompson, Kirkegaard & 37 Jahn, 1947 Telotroch formation in Scyphidia macropodia Davis, 1947 37 Telotroch formation in Scyphidia tholiformis Surber, 1943 38 PRECONJUGATION FISSION 38 Preconjugation fission in Rhabdosty/a verna/is Stokes, 1887 38 Preconjugation fission in Vorticella mierostome Ehrenberg, 1830 40 Preconjugation fission in Lagenophrys tattersalli Willis, 1942 40 Preconjugation fission in Scyphidia tholiformis Surber, 1943 41 CONJUGATION 42 Conjugation in Rhabdosty/a verna/is Stokes, 1887 42 Conjugation in Vorticella mierostome Ehrenberg, 1830 44 Conjugation in Urceo/aria synaptae Cuénot, 1891 47 Conjugation in Trichodina spheroidesi Pad nos & Nigrelli, 1942 52 Conjugation in Scyphidia ameiuri Thompson, Kirkegaard & Jahn, 1947 53 Conjugation in Scyphidia macropodia Davis, 1947 53 Conjugation in Scyphidia tholiformis Surber, 1943 55 Chapter 4 RESULTS: BINARY FISSION IN MANTOSCYPHIDIANS AND ELLOBIOPHRYIDS 93 Binary fission in Mantoscyphidia spadiceae Botes, Basson & 100 Van As, 2001 Binary fission in Ellobiophrya ma/icu/iformis Peters, Van As, Basson & 103 Van As, in press Chapter 5 TELOTROCH FORMATION IN MANTOSCYPHIDIANS AND ELLOBIOPHRYIDS 112 Telotroch formation in Mantoscyphidia branchi Van As, Basson & 112 Van As, 1998 and M. spadiceae Botes, Basson & Van As, 2001 Shape and surface structures 113 Size 113 Behaviour and swimming action 118 Telotroch formation 118 Method of attachment to substrate 119 Live observations and comparison with other species 119 Telotroch formation in Ellobiophrya ma/icu/iformis Peters, Van As, 120 Basson & Van As, in press Shape and surface structures 121 Size 121 Behaviour and swimming action 121 Telotroch formation 121 Comparison with other ellobiophryids 122 Method of attachment to substrate 123 Chapter 6 CONJUGATION IN MANTOSCYPHIDIANS AND ELLOBIOPHRYIDS 134 Conjugation in Mantoscyphidia branchi Van As, Basson & Van As, 134 1998 and M. midae Botes, Basson & Van As, 2001 Size 134 Appearance of peristomial regions 135 Attachment 135 FIRM ATTACHMENT AND ENTERING 135 PROGAMIC NUCLEAR DIVISIONS AND SHEDDING OF THE MICROCONJUGANT'S PELLICLE 136 SYNKARYON FORMATION 136 METAGAMIC (POST-ZYGOTIC) NUCLEAR DIVISIONS AND REORGANIZATIONAL FISSIONS 137 Conjugation in Ellobiophrya maliculiformis Peters, Van As, Basson & 137 Van As, in press Chapter 7 DISCUSSION 155 7.1 Molluscs and their scyphidiid peritrichs 156 Host adaptations to physical stress 157 Host/Symbiont Associations 158 Host/Scyphidiid peritrich Association 160 The effect of pollution 161 7.2 Binary fission 163 Mantoscyphidia 163 Ellobiophrya maliculiformis 168 7.3 Telotroch formation 170 Mantoscyphidia 170 Ellobiophrya maliculiformis 172 7.4 Conjugation 175 Mantoscyphidia 175 Ellobiophrya maliculiformis 177 7.5 FUTURE STUDIES 179 Chapter 8 REFERENCES 183 APPENDIX A TABLES AND RAW DATA 193 APPENDIX B TERMINOLOGY 206 APPENDIX C 213 To be submitted for publication in Acta Protozoologica: Peters, H., Van As, L..L., Basson, L. & Van As, J.G. In press. A new species of Ellobiophrya (Chatton & Lwoff, 1923) (Ciliophora: Peritrichia attached to Mantoscyphidia Jankowski, 1980 (Ciliophora: Peritrichia) species. ABSTRACT/OPSOMMING 214 ACKNOWLEDGMENTS 216 Chapter 1 Introduction , Chapter 1 Introduction 1 During the early 19th century a lot of progress was made in clarifying reproduction patterns and correlating cytological events with reproductive cycles in the kingdom Protozoa Goldfuss, 1818. During the first half of the zo" century various scientists studied the reproductive processes of peritrichs and made valuable contributions (Enriques 1907; Popoff 1908; Awerinzew 1912; Caullery & Mesnil 1915; Oilier 1928; Finley 1936,1939; Rosenberg 1940; Padnos & Nigrelli 1942; Finley 1943; Colwin 1944; Davis 1947; Finley 1952; Dobrzarïska 1961; Vávra 1961). Although thousands of recent papers exist on reproduction of ciliates, not much work has been done on epibiontic peritrichs (Lom personal communication)". Some of the most interesting reproductive phenomena are to be found in the subclass Peritrichia Stein, 1859. Binary fission, the formation of telotrochs, pre- conjugation fission and conjugation are well-established phenomena in this subclass (Finley 1952; Walker, Roberts & Usher 1986). Binary fission, budding and the formation of telotrochs are asexual methods of reproduction while pre-conjugation fission is the conjugant or sex differentiating fission, and conjugation is the sexual method of reproduction. The Aquatic Parasitology Research Group in the Department of Zoology and Entomology at the University of the Free State has been involved in studying parasites and symbionts of aquatic organisms since 1980. Most of their research has been devoted to freshwater organisms, but also included studies on intertidal species. Currently, most freshwater research in the Research Group forms part of the Okavango Fish Parasite Project. This project was initiated in 1997 and has already led to a number of scientific publications. I Prof. Jiri Lom, Institute of Parasitology, Academy of Sciences of the Czech Republic, Branisovska 31, 37005 Ceské Budejovice, Czech Republic. Chapter1 Introduction 2 Since 1994 the Foundation for Research Development (FRO), now referred to as the National Research Foundation (NRF), has been supporting their research project entitled: Intertidal Symbionts of the South African coast. This project falls within the realm of the Coastal Resources Program of the NRF. Within the context of this research program, two Ph.D. and six M.Sc. students have already completed their research on aspects of intertidal parasites and symbionts. Van As (1997) studied ciliophoran parasites of limpets (Patellogastropoda) and Smit (2000) studied the biology of gnathiid isopods and their role as vectors of fish blood parasites, both of these were Ph.D. studies. The M.Sc. works are that of Botha (1994) who studied ciliophoran symbionts of Oxystele Philippi, 1847 species; while Loubser (1994) studied the ciliophorans of intertidal fishes; Molatoli (1996) investigated the symbionts of red bait, Puyra stolonifera (Helier, 1878); Smit (1997) studied gnathiid isopods of intertidal fishes and Grobler (2000) investigated caligid fish parasites. The current author, Peters (néé Botes) studied scyphidiid peritrichs associated with Haliotis species (Botes 1999). These studies have led to the publication of various articles, congress proceedings and extended abstracts. Currently other M.Sc. projects are also being carried out and various Honours and Final year Zoology projects have also been completed within this program. Since the current study only deals with peritrichs only research concerning this will be listed. Full length publications (Basson & Van As 1992; Loubser, Van As & Basson 1995; Van As & Basson 1996; Van As, Basson & Van As 1998; Basson, Botha & Van As 1999; Van As, Van As & Basson 1999a; Van As & Van As 2000; Botes, Basson & Van As 2001a, Van As & Van As 2001; Peters, Van As, Basson & Van As in prep [see Appendix C)); congress proceedings (Van As, Van As & Basson 1995; Van As, Van As & Basson 1996a; Botes, Basson & Van As 1997; Van As, Basson & Van As 1997; Botes, Basson & Van As 1998; Van As, Van As & Basson 1998; Van As, Van As & Basson 1999b; Botes, Van As, Basson & Van As 2001, as well as published abstracts (Botha & Basson
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