Physical and Chemical Controls on the Abundance and Composition of Stream Microbial Mats from the Mcmurdo Dry Valleys, Antarctica
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PHYSICAL AND CHEMICAL CONTROLS ON THE ABUNDANCE AND COMPOSITION OF STREAM MICROBIAL MATS FROM THE MCMURDO DRY VALLEYS, ANTARCTICA by TYLER JOE KOHLER B.S., Kansas State University, 2007 M.S., University of Nebraska, 2010 A thesis submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of the requirement for the degree of Doctor of Philosophy Environmental Studies Program 2015 This thesis entitled: Physical and chemical controls on the abundance and composition of Dry Valley stream microbial mats written by Tyler Joe Kohler has been approved for the Environmental Studies Program Diane McKnight Patrick Kociolek Date The final copy of this thesis has been examined by the signatories, and we find that both the content and the form meet acceptable presentation standards of scholarly work in the above mentioned discipline. Kohler, Tyler Joe (Ph.D., Environmental Studies) Physical and chemical controls on the abundance and composition of stream microbial mats from the McMurdo Dry Valleys, Antarctica Thesis directed by Professor Diane M. McKnight ABSTRACT The McMurdo Dry Valleys of Antarctica are a cold, dry desert, yet perennial microbial mats are abundant in the ephemeral glacial meltwater streams that flow during austral summers. Three types of mats are present (orange, black, and green), and are primarily comprised of filamentous cyanobacteria, Nostoc, and chlorophytes, respectively. Mat types furthermore occupy distinct habitats within streams, utilizing the benthos, hyporheic zone, and water column, which expose them to different environmental conditions. Due to a lack of lateral inflows, allochthonous organic inputs, and negligible grazing activity, these streams are ideal for the controlled ecological study of microbial mats. Here, I investigated how mats will respond to physical disturbance, alterations in the hydrologic regime, and nutrient liberation from permafrost melt in the future. Specifically, I: 1) quantified and characterized the regrowth of mat biomass, community structure, and elemental stoichiometry after a scouring disturbance, 2) investigated how geomorphology and taxonomic identity influences the response of mat biomass to hydrologic regime in transects monitored over two decades, and 3) evaluated relationships between water chemistry and the elemental and isotopic composition of mat types over longitudinal and valley-wide gradients in Taylor Valley. I found that mats recovered ~20-50% of their biomass over the course of an austral summer following scour. Algal communities were significantly different in composition between disturbed and control treatments, but all samples naturally varied in species and elemental stoichiometry over the study period. When the long- term record of mat biomass was compared with hydrologic variables, stream channel mats (orange and green) had the greatest correlations, while marginal mats (black) showed weaker relationships with flow regime. Relationships also differed as a function of stream geomorphology, indicating the importance of substrata and gradient in conjunction with discharge. Lastly, mats showed unique elemental and isotopic compositions. Green and orange mats within the stream channel most reflected water column nutrient concentrations, while black mats showed significant nitrogen fixation. These results highlight the importance of taxonomic identity and habitat to modeling primary production here and elsewhere, and provide insight to how stream microbial mat communities are formed, maintained, and ultimately persist in an isolated polar desert. iii ACKNOWLEDGEMENTS There is no way to adequately thank all the people that have helped with these projects over the last five years, and it is similarly problematic to make a comprehensive list of everyone that I am indebted to. Please forgive me, for I have certainly left a lot of people out. For the rest, here is your “green dot.” First and foremost, I would like to thank my advisor, Diane McKnight. Not only did she support me even when doing things that made entirely no sense, but she can also make a spectacular strawberry-rhubarb pie. I would also like to thank my committee members, Pat Kociolek, Dan Liptzin, Jeb Barrett, and Sarah Spaulding for all their intellectual and emotional support over the last five years. Last but not least, thanks to my unofficial advisor Lee Stanish, for all of the emergency meetings and for leaving me a solid phycological path to follow. There is a special comradery between Dry Valley castaways with nothing to do on a weather day but play Settlers of Catan and Wizards. I would therefore like to give a shout out to my Antarctic family: Adrian Howkins, Jenny Erwin, Adam Lost-house-key, Emily Bernzott, Dave Van Horn, Alia Khan, Jon Denner, Garret Rue and his stupid trophy, Devin Castendyk, Aneliya Sakaeva, and Steven “Stinky Cheese” Crisp. A very special green dot goes to the big brother I never had, Chris Jaros, for teaching me all the stuff I probably should have already known. Thanks Chris. Green dots also go to honorary Asgard Rangers Kathi (and best office mate ever!) Hell, Kathy Welch, Thomas (Taco) Nylen, Amy Chiuchiolo, and Hilary Dugan. The Lake Hoare camp staff, Rae Spain and Kathy Schroeder, made sure I was well fed, clean, and on time for helicopters (93% success rate – good enough for a green dot). Thanks also to the McMurdo carpenters, PHI helicopter pilots, Crary Laboratory staff, and the awesome people of MacTown that made all the field campaigns successful. While this dissertation deals with algae in Antarctica, the bulk of the last half-decade was not spent there. In fact, it was spent in Boulder, Colorado, sometimes even away from INSTAAR, with some of the greatest people on Earth. That said, a very special thanks to John Berggren for being an incredible roommate, despite operating under what many would call challenging conditions. RIP Sev. Same goes for Bobby Moelter, and without his direction and encouragement, I would not have my own mug at the Walnut, nor would I be able to consistently out-fish John. To all the other Edge folks – Jessie, Katie, Matt, Theo, Tristan, Sarah, Mike, Megan, Alex, Margi – you guys are the best, thanks for being friends. For help with sample analyses, I would like to thank both the Kiowa Lab at the University of Colorado (Holly Hughes, Hanna Ding, Holly Shuss, Michael Brunetti) and Stefania Mambelli at the University of California Berkeley Center for Stable Isotope Biogeochemistry, both of whom rocked it. Thanks also to Samantha Weintraub and Tim Seastedt for allowing me to use their instruments, Eric Parrish for his help with maps, and Shelly Sommer for looking for papers that I promise exist. Steve Juggins and Dan Liptzin were indispensable in their statistical assistance. A very large, enthusiastic thank you goes to Chad Stoffel and Tara Hess for always fixing my computer and showing me how to make it stop sounding like a lawnmower (even if the solution was to stick a pencil in it), and to Penny Bates – sorry for all the migraines, and thanks for all you do for us. iv “Accurate” taxonomic identifications would not have been possible without opinions from Kateřina Kopalová, Jiři Komárek, Otakar Strunecký, Eci Jovanovska, Zlatko Levkov, Kalina Manoylov, Bart Van de Vijver, and Myriam de Haan. I would like to further thank the Distributed European School of Taxonomy, the Botanic Garden Meise, and Charles University in Prague for use of their facilities, and all the homies at Iowa Lakeside Laboratory for their invigorating discussions about algae and songs about Aphanizomenon. For nutrition during this hectic time, I would like to acknowledge King Soopers’ donuts (#2505), Big City Burrito, La Choza (at all locations), John and Theo’s home cooking, that Asian market on 28th street, The Walnut Brewery, Bumper Bars (but only the banana ones), Kate’s sack lunches, Swedish fish, and Bruce Vaughn for helping me achieve optimum daily caffeination with his coffee machine. I also owe Kate Kopalová and the Argentinians a big muchas gracias for introducing me to maté, which helped me back away somewhat from said coffee machine. Movement Climbing and Fitness chipped in with free hotdogs every July, and Tupac and Biggie provided the soundtrack. Financial support was provided as research assistantships by National Science Foundation Antarctic Organisms and Ecosystems Program (Award #0839020), the MCMLTER (OPP- 1115245), as well as numerous TA positions generously offered through Environmental Studies and under the supervision of Lisa Dilling (also an honorary committee member), Ryan Vachon, Max Boycoff, Dan Doak, and Jason Neff. Lastly, I would like to acknowledge waldeinsamkeit for keeping me off the streets, Brady Kohler for being an amazing brother and Yugioh opponent, Mom and Dad for having faith that one day I will have a real job (although I still don’t have a real job), and the mates back home that always have a PBR waiting. See ya Tuesday, -Tabasco Kid v CONTENTS CHAPTER I. ANTARCTIC STREAM ECOLOGY ......................................................1 Introduction .........................................................................................1 Arrangement of dissertation ..............................................................12 References .........................................................................................15 II. RECOVERY OF ANTARCTIC STREAM EPILITHON FROM SIMULATED SCOURING EVENTS ............................................................................22