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Diversity and Distribution of Epiphytic Diatoms

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Diversity and Distribution of Epiphytic Diatoms Palacký University Olomouc Faculty of Science Department of Botany DIVERSITY AND DISTRIBUTION OF EPIPHYTIC DIATOMS Ph.D. Thesis Mgr. Markéta Letáková Supervisor: Prof. RNDr. Aloisie Poulíčková, CSc. Olomouc 2018 1 ACKNOWLEDGEMENTS In the first place, I would like to thank Professor Aloisie Poulíčková for giving me the occasion to enter a world of science, which changed my view of many things, and for her kind supervision which was professional and friendly at the same time. Secondly, I wish to thank Dr. Marco Cantonati, for transferring to me his great enthusiasm for the work with diatoms and for teaching me the diatom morphology. I am grateful also to Doc. Petr Hašler and Dr. Markéta Fránková, for their various precious advices and inspiring cooperation, as well as to Dr. Nicola Angeli, for showing me his cool laboratory improvements which I have been using ever since. I would like to express my gratitude to all my colleagues of the Department of Botany for an inspiring and pleasant environment. Special thanks to Timothy Jones for the English corrections. Above all, I would like to thank to my family which has always been a great support for me. The research was funded by the Internal grant agency of Palacký University: PrF-2014-001, PrF-2015-001, PrF-2016-001 and PrF-2017-001. 2 DECLARATION I declare that I wrote this Ph.D. only by myself. All the sources used are included in References. The published results have been approved with the help of mentioned co-authors. 3rd January, 2018, Olomouc Mgr. Markéta Letáková 3 ABSTRACT This work is focused on one group of sessile diatoms that live attached to water plants – epiphytic diatoms. Many aspects of their life are unique and have not been explored enough yet, even though they may create considerable parts of primary production, and despite their importance in biomonitoring and paleolimnological reconstructions. The questions of substrate specificity, epiphytic diatom diversity, the range of species complexes within epiphytic diatom communities and the methods used for evaluation of the ecological status were investigated. The study of epiphytic diatoms was based on the diatom sampling and measuring the ecological parameters in the field, preparation of the permanent diatom slides, analysis of the species composition of the diatom communities and the statistical evaluation of acquired data. The aim of the work was to find the answers to the following question: i) Does the epiphytic diatom composition reflect the ecological parameters of the locality? ii) Does the type of macrophyte sampled influence the community composition? iii) What is the overall diversity of the epiphytic diatom communities? iv) What is the composition and frequency of the problematic species complexes? v) Do the unresolved species complexes reflect the state of water less precisely than other species do? vi) Are there any species complexes that if resolved could improve biomonitoring? Epiphytic diatoms reflected the ecological parameters of water in all investgated localities. The host-substrate preference was negligible in waters of higher trophy, in the lower trophy some significant specificity was found. Epiphytic assemblages have high overall diversity with considerable proportion of species complexes consisting of cryptic or semi- cryptic species. Although the pure genus level identification seemed appropriate for the routine monitoring, fine taxonomic resolution still shows sufficient variance related to the 4 environmental variable and increases the sharpness of classification. Species complexes showed lower sensitivity to change in phosphorus concentration than the rest of species. Some cryptic species have potential to improve bio-assessment models. Therefore resolving the species complexes is a crucial and urgent issue that would result also in better understanding the ecological state of waters. 5 ABSTRAKT (ČESKY) Tato disertační práce je zaměřena na jednu skupinu bentických rozsivek – na epifytické rozsivky, tj. na ty, které žijí přisedle na rostlinném substrátu. Mnoho aspektů jejich života není totiž stále dostatečně objasněno, přestože mohou tvořit velmi podstatnou část primární produkce a navzdory jejich důležitosti v biomonitoringu a paleolimnologických rekonstrukcích. Byla zkoumána substrátová specifita, diverzita epifytických rozsivek, rozsah druhových komplexů ve společenstvu epifytických rozsivek a také metody používané pro hodnocení ekologického stavu vod. Tato studie byla založena na vzorkování, měření ekologických parametrů v terénu, na přípravě trvalých rozsivkových preparátů, analýze druhového složení rozsivkových společenstev a na statistickém zpracování získaných dat. Cílem práce bylo najít odpovědi na následující otázky: i) Reflektuje složení epifytických rozsivek ekologické parametry lokality? ii) Ovlivňuje typ vodní rostliny složení společenstva epifytických rozsivek? iii) Jaká je celková diverzita společenstva epifytických rozsivek? iv) Jak velká část společenstva je tvořena problematickými druhovými komplexy, a jaké je jejich složení? v) Reflektují nerozlišené druhové komplexy stav vody méně přesně než jiné druhy? vi) Existují druhové komplexy, jejichž rozlišení by mohlo zkvalitnit biomonitoring? Epifytické rozsivky odpovídaly na ekologické parametry vody ve všech zkoumaných lokalitách. Substrátová specifita byla zanedbatelná ve vodách vyšší trofie, v čistém horském jezeře Valagola byla nalezena signifikantní substrátová specifita. Skupina epifytických rozsivek má celkově velkou druhovou bohatost a podstatná část je tvořena druhovými komplexy, tvořenými kryptickými a semikryptickými druhy. Ačkoliv se pouhé rodové určení zdá být naprosto dostačující pro rutinní monitoring, jemné taxonomické rozlišení stále vykazuje odchylku v souvislosti s proměnnými prostředí a zvyšuje přesnost klasifikace. 6 Nerozlišené druhové komplexy měly nízkou citlivost pro změny v koncentraci fosforu. Některé kryptické druhy mají potenciál zlepšit bioindikační modely. Proto rozlišení těchto druhových komplexů je zásadní a naléhavou záležitostí, která by vedla mimo jiné k lepšímu pochopení ekologického stavu vod. 7 CONTENT 1. Introduction ....................................................................................................................... 10 1.1 General diatom characteristics ........................................................................................ 10 1.2 Diatom age, diversity and ecological functions .............................................................. 10 1.3 Life strategies ................................................................................................................. 11 1.3.1 Planktonic diatoms ................................................................................................... 11 1.3.2 Benthic diatoms ....................................................................................................... 12 1.4 Epiphyton - Paper I ......................................................................................................... 14 2. Objectives of the dissertation ............................................................................................ 15 3. Methods ............................................................................................................................. 16 3.1 Material, sampling and preparation ................................................................................ 16 3.2 Measuring ecological parameters ................................................................................... 17 3.3 Statistical analysis ........................................................................................................... 17 4. Results ............................................................................................................................... 18 4.1 Paper II ........................................................................................................................... 19 4.2 Paper III .......................................................................................................................... 20 4.3 Paper IV .......................................................................................................................... 21 4.3 Paper V ........................................................................................................................... 22 5. Conclusions ....................................................................................................................... 23 5.1 Species richness and composition of epiphytic diatom communities ............................ 23 5.2 Epiphytic diatoms and ecological parameters ................................................................ 23 8 5.3 Substrate specificity problematics and implications to biomonitoring .......................... 24 5.4 Proportion of species complexes within epiphytic assemblages .................................... 25 5.5 Species complexes indicatory power and taxonomic resolution in biomonitoring ....... 26 5.6 Species complexes of epiphytic diatoms with potential to improve monitoring ............ 26 6. References ......................................................................................................................... 28 9 1. INTRODUCTION 1.1 General diatom characteristics Diatoms are unicellular autotrophic organisms, belonging into the group of Stramenopiles (Adl et al., 2012). Their typical feature is a silica cell wall – a frustule. They appear all over the world in various aquatic or wet terrestrial
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