Bangor University DOCTOR of PHILOSOPHY
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Bangor University DOCTOR OF PHILOSOPHY Studies on micro algal fine-structure, taxonomy and systematics : cryptophyceae and bacillariophyceae. Novarino, Gianfranco Award date: 1990 Link to publication General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal ? Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 02. Oct. 2021 Studies on microalgal fine-structure, taxonomy, and systematics: Cryptopbyceae and Bacillariopbyceae In Two Volumes Volume I (Text) ,, ý,ý *-ýýI Twl by Gianfranco Novarino, Dottore in Scienze Biologiche (Rom) A Thesis submitted to the University of Wales in candidature for the degree of Philosophiae Doctor University of Wales (Bangor) School of Ocean Sciences Marine Science Laboratories Menai Bridge, Isle of Anglesey, United Kingdom December 1990 Lýýic. JýýVt. BEST COPY AVAILABLE Acknowledge mehts Dr I. A. N. Lucas, who supervised this work, kindly provided his helpful guidance, sharing his knowledge and expertise with patience and concern, critically reading the manuscripts of papers on the Cryptophyceae, and supplying many starter cultures of the strains studied here. Mr T. B. B. Paddock and Dr D. G. Mann provided constructive criticism and valued suggestions on the manuscripts of the diatom papers. Drs P. Tett and D. K. Mills made it possible for me to perform contract phytoplankton analyses within the context of the NERC 'North Sea' research project 1988/89, and to include information derived therefrom in the present thesis. During the hot month of August, 1988, I was able to undertake some TEM preparation work at the Department of Evolutionary Biology, University of Siena, Italy. This was possible thanks to the kindness of Professors B. Baccetti (Siena), A. Burrini (Siena), and G. Ravagnan (Rome). Professors R. Gaddini de Benedetti (Rome) and G. B. Marini-Bettolo (Rome), were very helpful thanks to their general interest in my work. Professor G. Bazzichelli (Rome) and Dr N. Abdelahad (Rome), provided stimulating discussions during brief visits to Italy. Together with Dr L. Zucconi, Professor Bazzichelli also made diatom material from the Cesati Herbarium, Rome, available for study. Professor G. B. Fogg, FRS, kindly 'unearthed' precious information on the culture collection of R. W. Butcher, dating from the time it was being - iii - transferred from Burham-on-Crouch to Westfield College, London. Mr M. D. Budd, M. Phil., provided starter cultures of 'Rhodomonas' sp. We engaged in interesting discussions on biological philosophy, and (sometimes) managed to arrange lively musical sessions, which provided distraction and enjoyment. Within the context of the services provided by the EM Laboratory, School of Biological Sciences, University of Wales (Bangor), Mr D. A. Davies skilfully developed the photographic negatives taken with the Philips 301 and Kratos Corinth TEMs. Financial assistance was provided by Consiglio Nazionale delle Ricerche, Italy, in the form of a special contribution 'Altri Interventi' from September 1988 to March 1989; and from Accademia Nazionale dei Lincei, Rome, jointly with the Royal Society, London, in the form of an award from April 1990 to July 1990, within the context of the European Science Exchange Programme. I should like to pay a tribute to Sir Frederick Banting and Charles Best, whose pioneering work on the isolation of insulin in the 1920s eventually made it possible for countless insulin- dependent diabetics to enjoy life much in the same way as non-diabetics. Last, but not least, I should like to thank my loved ones, whose concern, support, and affection, made this thesis possible in the first place. To all of them, and especially Alfredo and Nica, this work is dedicated with gratitude. - iv - Abstract Part I deals with the fine-structure of 25 marine and 3 freshwater strains of cryptomonads, and 2 species observed in fixed marine plankton samples. Emphasis is placed on scanning electron microscope observations, which yield valuable taxonomic data, providing that cell shrinkage (up to about 25 % linear), and other undesirable effects of preparation schedules, are taken into account. The fine-structural information is used for judging the taxonomic value of some cell features. Characters having taxonomic significance at various levels include the cell shape, periplast features (especially the shape and size of the periplast areas), chloroplast number, nucleomorph position, presence of a 'tail', and the presence of a true, non-artifactual furrow on the ventral cell face. A new classification system is proposed. This allows for the existence of three orders, four families, seven pigmented genera, and two colourless ones. Most diagnosis or descriptions of taxa above the generic level are based on only one character. This should make the system capable of accomodating new taxa with little difficulty. The following new taxa are described: Rbinomonas reticulata comb. nov., R. et stat. nov., R. .reticulata var. atrorosea comb. reticulata var. compresses var. nov., R. reticulate var. eleniana var. nov., Proteomonas pseudobaltica comb. nov., P. pseudobal tics var. 1 eonardiana var. nov., Pyrenomonas sahne var. curvata var. nov., Pyrenomonadales ord. nov., Pyrenomonadaceee fam. nov., and Cryptomonadales ord. nov. - Part II deals with the diatom genus Mastoglola. An update of the taxa assigned to this genus is given, together with a 'resemblance list' useful for identifying taxa not included in the main reference work for the genus. The frustule architecture of Mastoglola smitbii is examined with particular reference to the valvocopula, especially the features involved in its integration with the valve. - vi - Fore word Whether the facts of nature are only awaiting to be discovered, or whether they are revealed to us with parsimony, is a matter which may be adequately discussed by philosophers alone. In the case of the cryptomonads, however, it is unnecessary to be versed in philosophy for agreeing fully with the second point of view. These organisms have been studied for more than a century and a half, and yet remain an intriguing and mysterious group of algae, protozoans, protists, protoctists, chromists, or ... M. The first studies on their fine-structure brought about the feeling that their secrets would soon be unlocked. It appears that this promise has not yet been fulfilled: the cryptomonads, which have cast new light on the origin and evolution of the eukaryotic cell, have been extremely reticent about themselves. Have they given all that they can give, or can they tell us more? Could it not be that the notorious difficulties of their taxonomy and systematics are due, at least in part, to some kind of myth? It is hoped that Part I of this work may provide a clue, however modest, towards the answer. The diatoms are probably the most intensively studied group of microalgae, and yet they still raise numerous questions: for instances, the process of identifying species within the large genera; the structure of the frustule, especially the architecture and integration of the various parts; and the phylogenetic relations between genera. It is hoped that Part II of this work may contribute some information of interest. - vii - Third Party material excluded from digitalized copy. Please refer to original text to see this material. - viii - CC n- ents Declarations ii Acknowledgements Iii Abstract v Foreword vii Some philosophical considerations: general introduction to this thesis 1 Part I: Cryptophyceae Chapter 1- Introduction to Part I <Cryptophyceae)8 Chapter 2- Material and methods 17 Chapter 3- Observations on Hemiselmis rufescens and Hemiselmis virescens, with particular reference to the periplast 29 Chapter 4- Observations on new varieties and combinations in the genus Rbinomonas, with comments on the genera Pyrenomonas and Rbodomonas 38 Chapter 5- Observations on Proteomonas pseudobalti- ca comb. nov. and Proteomonas pseudobaltica var. leonardiana var. nov., with comments on a freshwater Cryptomonas 599 ' Chapter 6-A brief scanning electron microscope survey of marine Cryptophyceae, with comments on a freshwater form, cell shrinkage, and some effects of fixation schedules 69 - ix - Chapter 7- Observations on some species of Cbroomonas 93 Chapter 8- Some proposals for a new classification system of the Cryptophyceae 104 Chapter 9- Discussion to Part 1 124 Part II: Bacillariophyceae and general conclusions II Chapter 10 - Introduction to Part (Bacillariophyceae) 130 Chapter 11 - An update of the taxa of the genus Nastogloia, with a 'resemblance li st' for the more recently described ones 133 Chapter 12 - Observations on the frustule architecture of Xastogloia smitbii 136 Chapter 13 - Discussion to Part II 148 Chapter 14 - Mare philosophical con siderations: general conclusions to this thesis 153 Appendix -A database of selected culture strains of Cryptophyceae 155 References 214 - Some