Systematics of Coccal Green Algae of the Classes Chlorophyceae and Trebouxiophyceae
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School of Doctoral Studies in Biological Sciences University of South Bohemia in České Budějovice Faculty of Science SYSTEMATICS OF COCCAL GREEN ALGAE OF THE CLASSES CHLOROPHYCEAE AND TREBOUXIOPHYCEAE Ph.D. Thesis Mgr. Lenka Štenclová Supervisor: Doc. RNDr. Jan Kaštovský, Ph.D. University of South Bohemia in České Budějovice České Budějovice 2020 This thesis should be cited as: Štenclová L., 2020: Systematics of coccal green algae of the classes Chlorophyceae and Trebouxiophyceae. Ph.D. Thesis Series, No. 20. University of South Bohemia, Faculty of Science, School of Doctoral Studies in Biological Sciences, České Budějovice, Czech Republic, 239 pp. Annotation Aim of the review part is to summarize a current situation in the systematics of the green coccal algae, which were traditionally assembled in only one order: Chlorococcales. Their distribution into the lower taxonomical unites (suborders, families, subfamilies, genera) was based on the classic morphological criteria as shape of the cell and characteristics of the colony. Introduction of molecular methods caused radical changes in our insight to the system of green (not only coccal) algae and green coccal algae were redistributed in two of newly described classes: Chlorophyceae a Trebouxiophyceae. Representatives of individual morphologically delimited families, subfamilies and even genera and species were commonly split in several lineages, often in both of mentioned classes. For the practical part, was chosen two problematical groups of green coccal algae: family Oocystaceae and family Scenedesmaceae - specifically its subfamily Crucigenioideae, which were revised using polyphasic approach. Based on the molecular phylogeny, relevance of some old traditional morphological traits was reevaluated and replaced by newly defined significant characteristics. Declaration [in Czech] Prohlašuji, že svoji disertační práci jsem vypracovala samostatně pouze s použitím pramenů a literatury uvedených v seznamu citované literatury. Prohlašuji, že v souladu s § 47b zákona č. 111/1998 Sb. v platném zněni souhlasím se zveřejněním své disertační práce, a to v nezkrácené podobě elektronickou cestou ve veřejně přístupné časti databáze STAG provozované Jihočeskou univerzitou v Českých Budějovicích na jejich internetových stránkách, a to se zachováním mého autorského práva k odevzdanému textu teto kvalifikační práce. Souhlasím dále s tím, aby toutéž elektronickou cestou byly v souladu s uvedeným ustanovením zákona č. 111/1998 Sb. zveřejněny posudky školitele a oponentů práce i záznam o průběhu a výsledku obhajoby kvalifikační práce. Rovněž souhlasím s porovnáním textu mé kvalifikační práce s databází kvalifikačních prací Theses.cz provozovanou Národním registrem vysokoškolských kvalifikačních prací a systémem na odhalovaní plagiátů. České Budějovice 3. 11. 2020 Lenka Štenclová This thesis originated from Faculty of Science, University of South Bohemia, supporting doctoral studies in the Botany study program. Financial support: Financial support came from the budged of the Department of Botany (RVO) and partly from Assumption College Faculty Development Grant. Acknowledgements: On the very first place I thank to my supervisor Hanys for not stopping believing at this project, for his support and great ideas. I owe my thanks to Karolina Fučíková as well, for her friendly cooperation on Assumption college. I thank my colleagues and friends at the same time from České Budějovice for valuable advices and all their funny talks that this is thesis full of small green balls looking all alike. My special thanks belong to my parents for their kind support. List of papers and author’s contribution The thesis is based on the following papers (listed according to the thematic chapters): Štenclová, L., Fučíková, K., Kaštovský, J. and Pažoutová, M. (2017) Molecular and morphological delimitation and generic classification of the family Oocystaceae (Trebouxiophyceae, Chlorophyta). Journal of Phycology. 53(6):1263-1282. (IF 2017 = 3.000) Lenka Štenclová cultivated strains, performed molecular methods and molecular phylogeny, morphological and ultrastructural (TEM) observations, and was main author of the manuscript. Silva, T. G., Štenclová, L., Archanjo, N.C.P. and Bagatini, C. L. Revised phylogenetic position of genus Nephrocytium Nägeli (Sphaeropleales, Chlorophyceae), with description of Nephrocytiaceae fam. nov. and Nephrocytium vieirae sp. nov. (Manuscript) Lenka Štenclová cultivated one strain, performed molecular methods, morphological and ultrastructural (TEM) observations, and was second author of the manuscript. Štenclová, L. Distribution of the Crucigenioid algae inside the classes Chlorophyceae and Trebouxiophyceae. (Manuscript) Lenka Štenclová cultivated strains, performed molecular methods and molecular phylogeny, morphological observations, and was only author of the manuscript. Štenclová, L. and Fučíková, K. (2019) Dispora speciosa, a new addition to the genus Parallela and the first coccoid member of the family Microsporaceae. Phytotaxa. 419(1):63-76. DOI:10.11646/phytotaxa.419.1.4 (IF 2018/19 = 1.168) Lenka Štenclová cultivated strain, performed molecular methods, morphological and ultrastructural observations, and was main author of the manuscript. Contents Chapter 1: Introduction: Situation in current systematics of green coccal algae............... 1 1.1 Traditional morphological system .......................................................................... 2 1.2. Ultrastructural research challenging the morphological system ............................ 5 1.3. Modern system of green algae based on the polyphasic approach ........................ 7 1.4. Coccal algae inside classes Trebouxiophyceae a Chlorophyceae........................ 10 1.5. Ongoing problems in current systematics of green coccal algae ......................... 15 1.6. Phylogenetic methods as the backbone for modern systematics ......................... 18 1.7. Aims of the thesis ................................................................................................ 20 1.8. Results ................................................................................................................. 21 1.8.1 Molecular and morphological delimitation and generic classification of the family Oocystaceae ........................................................................................ 22 1.8.2. Revised phylogenetic position of genus Nephrocytium Nägeli (Sphaeropleales, Chlorophyceae), with description of Nephrocytiaceae fam. nov. and Nephrocytium vieirae sp. nov. ............................................................... 23 1.8.3. Distribution of Crucigenioid algae in classes Chlorophyceae & Trebouxiophyceae ................................................................................................ 24 1.8.4 Dispora speciosa, a new addition to the genus Parallela and the first coccoid member of the family Microsporaceae ................................................. 26 1.9. Conclusion ........................................................................................................... 26 1.10. Following problems ........................................................................................... 27 1.10.1. Delimitation and species concept of the genus Oocystis ........................ 27 1.10.2. Phylogenetic position of Juraniella javorkae and new insides in the phylogeny of the genus Kirchneriella ................................................................. 28 1.10.3. Characterizing the chloroplast and mitochondrial genomes of a microscopic alga Oonephris obesa (Chlorophyceae) ......................................... 28 1.11. References ......................................................................................................... 30 Chapter 2 Molecular and morphological delimitation and generic classification of the family Oocystaceae (Trebouxiophyceae, Chlorophyta) ............................................ 45 Chapter 3 Revised phylogenetic position of genus Nephrocytium Nägeli (Sphaeropleales, Chlorophyceae), with description of Nephrocytiaceae fam. nov. and Nephrocytium vieirae sp. nov. ....................................................................................... 103 Chapter 4 Distribution of Crucigenioid algae in classes Chlorophyceae & Trebouxiophyceae ......................................................................................................... 107 Chapter 5 Dispora speciosa, a new addition to the genus Parallela and the first coccoid member of the family Microsporaceae ............................................................ 111 Authors CV ……………………………………………………………………….145 Chapter 1: Introduction: Situation in current systematics of green coccal algae Pröschold and Leliaert 2007 1 Chapter 1: Situation in current systematics of green coccal algae 1.1 Traditional morphological system Coccal thallus is the simplest organization of the algal body because the entire organism consists of simply one cell. Coccal thallus covered by visible mucilage wrap is called palmeloid or also capsal. Both types can agglomerate into regular or irregular colonies and coenobia. All the coccal and palmeloid green algae were, according to the traditional morphological system summarized in the last comprehensive morphological monograph by Komárek and Fott (1983), assembled in a single order Chlorococcales, just on the basis of the common body organization Komárek and Fott (1983), expecting its one time evolution from a green flagellate ancestor common for