Terrestrial Influences on the Macroinvertebrate Biodiversity of Temporary Wetlands
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TERRESTRIAL INFLUENCES ON THE MACROINVERTEBRATE BIODIVERSITY OF TEMPORARY WETLANDS Michael A. Plenzler A Dissertation Submitted to the Graduate College of Bowling Green State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY December 2012 Committee: Dr. Helen Michaels, Advisor Dr. Enrique Gomezdelcampo Graduate Faculty Representative Dr. Jeff Miner Dr. Karen Root Dr. Amy Downing ! ii! ABSTRACT Dr. Helen J. Michaels, Advisor Vernal pools are temporary wetlands and local-scale biodiversity hot spots for a variety of amphibians, macroinvertebrates, and plants because their seasonal drying prevents the establishment of predatory fish populations. Vernal pools are often of conservation concern because of the amphibian populations; however, the emphasis on these organisms often eclipses the macroinvertebrates, which are important predators, prey, and nutrient cyclers in wetlands and the surrounding habitat. Hydroperiod and water chemistry are thought to be the primary regulators of vernal pool macroinvertebrates, but the surrounding habitat also affects these organisms. Specifically, canopy cover and forest composition can alter the autochthonous and allochthonous carbon sources for wetland food webs. My research objectives were to understand how variations in these factors affect macroinvertebrate diversity and community composition. In 2009, I conducted a field survey of fifteen vernal pools that varied in area, depth, hydroperiod, canopy and surrounding land use. I measured several habitat conditions, assessed the biotic communities of these wetlands, and found that canopy cover influenced bottom-up productivity and macroinvertebrate diversity. I used the results of this study to determine how known macroinvertebrate communities respond to variation in canopy cover in mesocosm wetlands. The low canopy treatments sustained the highest macroinvertebrate abundance, family richness, and Shannon diversity, as greater algal productivity increased resources available to support the macroinvertebrate communities. I conducted a second mesocosm experiment to explore how variation in canopy cover (low or high) and litter species (oak, maple, or a mixture of the two litters) affect vernal pool macroinvertebrates. Macroinvertebrate abundance and family richness were greater under low canopy and with oak litter, but the effects of canopy often depended on ! iii! litter species. Canopy limits the amount of light that reaches a wetland, but litter and its impact on dissolved organic carbon (DOC) limit light transmittance and producer communities. Because forest composition and structure are changing from anthropogenic activities, these results suggest they will also alter the food webs of vernal pools. However, as some organisms utilize pools considered to be low quality for macroinvertebrates, vernal pools should be protected in a variety of habitat contexts to preserve regional species diversity. ! iv! ACKNOWLEDGEMENTS I once read that a PhD isn’t a statement of intelligence, but rather one of endurance. In my experience, it’s also one of luck and being surrounded by the right people at the right time. I can’t express the number of trying times I’ve encountered on this journey, but I’ve always been amazed at the number of people standing behind me throughout this process. To that end, there’s no shortage of people deserving some sort of recognition here (and I’ll take my time doing it). First off, my committee. Thank you for helping me take an off-the-cuff observation about vernal pools and turn it into this project. It’s my sincere hope that you see your fingerprints all over this work, too. You have all pushed me further than I ever anticipated going—or sometimes wanted to go. I can’t honestly remember why I chose to do this degree, but I do feel equipped to go out and change some corner of the world for the better. Dr. Enrique Gomezdelcampo was a fantastic outside referee who was equal parts informed contributor to this work and a willing combatant with an uppity stranger causing a stir at my conference talk. Dr. Jeff Miner pulled me into graduate school. I had no idea what I was doing then, but I have a better idea of what I’m doing now thanks to him. His keen understanding of aquatic ecology was instrumental in designing and interpreting the following experiments. By some stroke of luck, Dr. Amy Downing was invited to give a seminar and helped launch the mesocosm projects. She’s been a fantastic addition to this work and I suspect she’s a Swiss Army knife of statistical techniques, too, as many of the analyses in this dissertation are modeled from her papers. Dr. Karen Root’s emphasis on scale and conservation underpins much of my thinking on the need for vernal pool preservation. It’s through my conversations with her that I’ve come to think about these wetlands as metacommunities, where connections between them are critical and that habitat quality is organism-subjective. Finally, in one last act of Stockholm syndrome, my ! v! advisor Dr. Helen Michaels, who essentially allowed me to run free with a project that was a complete turn from anything that had previously gone on in the lab. I have a feeling I’ll be asking people, “What’s your question?” thirty years from now. It would feel glib to wax on about being the scientist I am now because of the years as her student, but it’s true. It’s odd thinking about the lack of Bowling Green in the next stage of my life, but as a testament to our history, I do plan on dropping in on her unannounced beyond publishing this work. To my dear lab mates…it will be an incredibly weird day when I walk into an office and not see all of your faces or trip over all of your chairs on the way to my desk. Between Ryan Walsh and Jake Meier, it’s been an odd marriage over the years and with the number of off-the- wall things that we’ve said and done, I think we’d have a hit radio show. Jen Shimola has been kind enough to not alert the authorities or psychologists about us. With everyone trying to finish up at once, we’ve probably recently carped at each other more than was necessary. I’ll miss it no less. Expect random phone calls in the future and countless more baseless thoughts about…well, anything goes. The lab and summer of 2012 were improved by the addition of four undergraduates. I’m sure you all have names, but I never really cared to learn any of them (I’m just kidding, calm down). Paige Arnold, Alyssa Dietz, Jake Sublett, and Alex Hill—at the very least, you were all a wonderful infusion of energy and ideas into the lab. If there was any crowd I had to be surrounded by during this process, I’m glad it was this one. None of this work would be possible without help from Dave Bonner (chapter one), Adam Meeker and Dante D’Avello (chapter two), and Nate Schweitzer (chapter three). I mean that. Thanks for your volunteer labor and the oft-interesting conversations in the field. I can’t emphasize enough just how important your contributions were to this project. Don Schmenk (Maumee State Forest), Gary Haase and Steve Woods (Kitty Todd), Chris Smalley (Wood ! vi! County Parks) and Karen Menard (Toledo Metroparks) were critical for helping me locate vernal pools and granting permit access to them. The Ohio Biological Survey, Graduate Student Senate, and BGSU Department of Biological Sciences Oman Fellowship provided highly appreciated funding for chapters two and three. Many, many, many thanks to all of you. There are a number of other people who helped me get up the mountain in some way, shape, or form. First off, my family—especially my parents and sister. You guys provided no end of support. It was incredible and I find myself at a loss for words to explain it. Whether it was a place to run away to or a free meal when I had little time to feed myself, it all added up. I don’t know if I’ll ever find a way to adequately express my gratitude, but I’ll be able to start soon by repaying the money I borrowed to help keep the lights on during this whole experience. For my friends in Toledo, Cleveland, Cincinnati, Columbus, Milwaukee, Chicago, San Francisco, Pittsburgh, Houston, Boston, Arkansas, and on, and on, and on. Thank you. Thank you for your encouragement, the random visits back to BG, and your infinite patience with my frequent existential crises and other grad school related sacrifices. I plan to make good on my promises to visit you all and with a little bit of luck, might just end up employed near you, too. You guys rock. I’d also like to give a special thanks to a number of other BG people. Dr. Rex Lowe provided assistance with identifying algae samples and always had an offbeat story about life as a scientist or BG resident. Dr. Dan Pavuk helped identify bugs and was always good for a friendly conversation. The McKay/Bullerjahn lab was kind enough to provide equipment and space for chlorophyll fluorometry and Ben Beall conducted some flow cytometry that ultimately confirmed my water samples were too old and not properly preserved for such a thing. Bob Midden and Sarah Jindra provided invaluable experience and help for analyzing nutrient content ! vii! in the water samples. Frank Schemenauer and the greenhouse were the perfect getaway when I got too cagey in the office. A lot of respect is also directed towards the Root and Miner labs. You always showed up to my talks, but were also fun traveling companions and, in a pinch, free therapy for the rigors of grad school.