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Open Mtaylor Finalthesis.Pdf THE PENNSYLVANIA STATE UNIVERSITY SCHREYER HONORS COLLEGE DEPARTMENT OF BIOLOGY DIVERSITY OF SYMBIOTIC DINOFLAGELLATES IN NORTHERN LATITUDE CORALS MADISON TAYLOR SPRING 2018 A thesis submitted in partial fulfillment of the requirements for a baccalaureate degree in Biology with honors in Biology Reviewed and approved* by the following: Todd LaJeunesse Associate Professor of Biology Thesis Supervisor James Marden Professor of Biology and Associate Director, Huck Institutes of the Life Sciences Honors Adviser * Signatures are on file in the Schreyer Honors College. i Abstract Reef building corals are widespread throughout the tropics and subtropics; however, some species exist at high temperate latitudes. Temperate conditions such as high seasonality in temperature and turbidity impact coral physiology and likely affect the endosymbiotic dinoflagellates (Symbiodinium) that live in their host tissues. The identities of endosymbionts from corals at extreme northern latitudes are poorly characterized, therefore we used molecular genetic approaches to examine Symbiodinium collected across a broad host taxonomic range around Honshu Island, Japan. We observed patterns of high specificity between coral and symbiont as well as a few generalist symbionts. In addition, our data provide evidence that many high latitude Symbiodinium species are new species well adapted to the seasonal fluctuations of temperate environments like Japan. These findings suggest a long-term co-evolution of symbionts that are highly adapted to the intracellular environment of particular coral species at high latitudes. This research improves our understanding of Symbiodinium diversity in high latitude coral communities and begins to elucidate to what extent this diversity is unique relative to the symbionts that thrive in coral populations at lower latitudes. Moreover, characterizing these symbioses is important for the future of coral reef conservation by establishing a baseline of biodiversity at northern latitudes, as corals may find protection from the effects of climate change by colonizing at northern latitudes like Japan. ii Table of Contents List of Figures .............................................................................................................. iii List of Tables ............................................................................................................... iv Acknowledgements ...................................................................................................... v Introduction .................................................................................................................. 1 The significance of coral-algal symbioses ....................................................................... 1 Defining Species of Symbiodinium .................................................................................. 3 Patterns of Symbiodinium evolution ............................................................................... 6 Symbiodinium distribution and ecology .......................................................................... 7 High vs. low latitude: the importance of studying symbioses at extremes ....................... 8 Investigating the diversity of Symbiodinium in northern latitude corals ........................ 11 Materials and Methods ................................................................................................. 13 Sample collection and preservation ................................................................................. 13 DNA extraction and Polymerase Chain Reaction (PCR) ................................................. 14 Denaturing Gradient Gel Electrophoresis (DGGE) and fingerprinting .......................... 15 Sanger sequencing and phylogenetic analysis ................................................................. 16 Coral taxonomy ................................................................................................................ 17 Results. ......................................................................................................................... 19 Symbiont diversity quantified with genetic evidence ........................................................ 19 DGGE analysis of Symbiodinium common to Acropora ................................................. 19 Ecological distribution of symbiont diversity................................................................... 21 Host diversity and phylogeny ........................................................................................... 24 Discussion .................................................................................................................... 26 High latitude lineages of Symbiodinium ......................................................................... 26 Ecological niche and host specificity ............................................................................... 28 Generalist species ............................................................................................................ 29 Effect of climate change on high latitude coral symbioses .............................................. 30 Limitations and further research ..................................................................................... 31 Supplemental Table ..................................................................................................... 33 Supplemental Figure .................................................................................................... 36 Bibliography ................................................................................................................ 37 iii List of Figures Figure 1. The host-symbiont relationship between coral and Symbiodinium .......................... 3 Figure 2. Collection sites on Honshu Island, Japan ................................................................. 14 Figure 3. Phylogeny of Symbiodinium collected from Japan based on the ribosomal marker LSU20 Figure 4. Phylogeny of Symbiodinium collected from Japan based on the mitochondrial marker cob .................................................................................................................................... 21 Figure 5. Phylogeny of Symbiodinium collected from Japan based on the chloroplast marker cp23s ................................................................................................................................ 22 Figure 6. Phylogeny of coral host families and their associated symbiont types ..................... 25 Figure 7. DGGE ITS2 results containing coral hosts Acropora and Montipora ...................... 36 iv List of Tables Table 1. List of sampled corals from Japan, their genus, species, and the number of specimens collected ........................................................................................................................... 33 v Acknowledgements “We just don’t get to be competent human beings without a lot of different investments from others” –Mister Rogers I have had so many incredible people supporting me through the thesis process. First, I’d like to express my gratitude to the entire LaJeunesse Lab Team for three years of incredible opportunities to learn what hands-on science looks and feels like. Todd, thank you for guiding me from the basics to where I am now, with a lot of memorable conversations in between. To Daniel, for being a smart friend and fellow marine scientist. Thanks to Kira, Hannah, and Allison for being boss women in science. I couldn’t have had better role models to experiment and do science with. Kira, thank you for being an inspiration both in the lab and out on the trails. Hannah, thanks for sharing office space with me all of the Fall semester and for always pulling my samples from the PCR machine. Last but not least, I’d like to express my endless gratitude to Allison for her patience, guidance, and mentorship over the years. Thank you for fielding my many questions, the FaceTime calls, shared laughs and honest talks about where we’re both going next. Thank you also to Dr. Marden for being a fantastic resource and honors advisor. Our trip to Costa Rica will forever be a huge influence on my future career; thank you for showing me how challenging yet rewarding fieldwork can be. Lastly, thank you to the Schreyer Honors College for academic support, thesis boot camps, and the opportunity to culminate my research in a body of work that I am proud of. 1 Introduction The significance of coral-algal symbioses Coral ecosystems are home to a wide array of marine organisms, earning their name of the “rainforests of the sea”. They provide a habitat and nutrition for fish, invertebrates, marine mammals, and plants, among other organisms. Coral reefs are also critical to aspects of human society: tourism, coastal and island community cultures, food security, and the conservation of biodiversity. Coral colonies are the foundation of reef ecosystems and if harmed, numerous other species and aspects of human society suffer as well. Currently, coral populations are threatened by environmental degradation. Climate change, ocean warming and acidification, pollution, and unsustainable resource management are putting stress on these animals and their symbiotic partners. Some of these threats cause massive bleaching events and the eventual death of reefs. Therefore, there is a pressing need to study corals, in order to inform future conservation practices. Coral reef ecosystems are widespread across the planet. They
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