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Thesis Filing Form University of Nevada, Reno The Dynamics of Mysis diluviana and Other Zooplankton in Three Oligotrophic Lakes A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in Biology by Zachary Bess Dr. Sudeep Chandra/Thesis Advisor August 2020 Copyright by Zachary A. Bess 2020 All Rights Reserved THE GRADUATE SCHOOL We recommend that the thesis prepared under our supervision by entitled be accepted in partial fulfillment of the requirements for the degree of Advisor Committee Member Committee Member Graduate School Representative David W. Zeh, Ph.D., Dean Graduate School i Abstract Mysids (Mysis diluviana) have been introduced to a number of temperate lakes to provide food to salmonids for recreational fishing. In many of these lakes, these mysids have changed the native zooplankton communities through predation. Mysids were introduced to Donner Lake, Fallen Leaf Lake, and Lake Tahoe in the mid-1960s and have changed the native zooplankton community of Lake Tahoe profoundly. We conducted mesocosm experiments to evaluate the effects of two native zooplankton taxa (Daphnia spp. and Epischura nevadensis) and juvenile and adult mysids on ecosystem function in Lake Tahoe and its more productive embayment, Emerald Bay. The results of these experiments indicate that these zooplankton play significantly different roles in the Emerald Bay ecosystem, but not in Lake Tahoe proper. This suggests that these zooplankton may play a larger role in shaping the ecosystem characteristics of the water column in Lake Tahoe if cultural eutrophication should eventually elevate Lake Tahoe’s trophic state to that of Emerald Bay’s. Additionally, we measured the environmental factors that influence mysid growth rates in Lake Tahoe. We found that adult growth rates significantly correlate with the depth of winter mixing, but juvenile growth rates significantly correlate with the mean summer. We also measured the pelagic reliance, trophic position, and carbon sources of mysids in Donner Lake, Fallen Leaf Lake, and Lake Tahoe and evaluated the role of wind-dispersed pollen in supporting mysid energetics. These analyses indicate that a variety of factors influence mysid production in these lakes, and the importance of the factors differs between the lakes. ii Acknowledgements I would like to thank my graduate advisor, Dr. Sudeep Chandra, for providing guidance, inspiration, funding, assurance, and a lab to work in. His tremendously broad knowledge of and passion for aquatic ecology have been evident and invaluable. He fostered an environment in which I was able to learn about many aspects of aquatic and conservation ecology beyond just what was relevant for this thesis. Additionally, the get- togethers at his home (complete with tandoori chicken and musical instruments) were a great way to take a break from the work of this thesis. The other members of my committee (Dr. Alan Heyvaert, Dr. Steve Sadro, and Dr. Zeb Hogan) honed the design of the experiments and provided edits to this thesis, and I am thankful for their guidance as well. Two of Dr. Chandra’s former students significantly helped to smooth my path. Karly Feher was an excellent mentor, especially in the early stages of working in Dr. Chandra’s lab. By assisting her at Castle Lake, I learned many of the methods that I have used in this thesis. She was also eager to offer her help when I needed it. Dr. Tim Caldwell helped to brainstorm ideas with me that made their way into both chapters of this thesis. He also offered his expertise in all things related to mysids. The other members of Dr. Chandra’s lab also made for spectacular collaborators. They include Erin Suenaga, Dr. Suzanne Kelson, Emily Carlson, Emily Carlson, Dr. Ed Krynak, Dr. Flavia Tromboni, Dr. Facundo Scordo, Dr. Emanuele Ziaco, James Simmons, Josh Culpepper, Teresa Campbell, Elizabeth Everest, and Loren Secor. Bonnie Teglas patiently and persistently managed the funds and expenses that supported this work. iii Several others were instrumental in helping me, often in ways that I did not initially realize and fully appreciate. In her graduate course at UNR, Dr. Mae Gustin provided me with advice when developing the project and helped me to develop a critical eye when reviewing others’ works. Likewise, another UNR professor, Dr. Kevin Shoemaker, helped me with improving my understanding of computer coding and statistics and even made recommendations for the tests employed in this thesis. Dr. Jim Hobbs of CA Department of Fish & Wildlife and Dr. Malte Willmes of UC Davis gave me my first exposure to the utility of stable isotope ecology. Dr. Peter Moyle of UC Davis inspired me through his applications of aquatic biology to conservation ethics. Brant Allen, Katie Senft, and Brandon Berry of the UC Davis Tahoe Environmental Research Center offered training and advice in working at Lake Tahoe and provided samples that were analyzed in Chapter 2. I thank my family for providing support while completing this endeavor. iv Table of Contents General Introduction………………………………………………………………………1 General Introduction References………………………………………………….4 Chapter 1. Zooplankton influences on phytoplankton, water clarity, and nutrients in Lake Tahoe………………………………………………………………………………………6 Abstract……………………………………………………………………………7 Introduction………………………………………………………………………..8 Materials & Methods…………………………………………………………….11 Results……………………………………………………………………………17 Discussion………………………………………………………………………..19 References………….…...………………………………………………………..25 Tables…………………………………………………………………………….32 Figures……………………………………………………………………………36 Supplemental Table……………………………………………………………...43 Chapter 2. Factors Influencing the Production of Mysis diluviana in Three Oligotrophic Lakes……………………………………………………………………………………..44 Abstract…………………………………………………………………………..45 Introduction………………………………………………………………………46 Methods…………………………………………………………………………..48 Results……………………………………………………………………………55 Discussion………………………………………………………………………..57 References………………………………………………………………………..64 Table……………………………………………………………………………..72 Figures……………………………………………………………………………73 Supplemental Tables……………………………………………………………..80 General Summary of Findings…….……………………………………………………..93 General Summary References……………………………………………………95 v List of Tables Chapter 1 1. Secchi depth, chlorophyll a concentrations, and SRP concentrations for Lake Tahoe and Emerald Bay. Lake Tahoe chlorophyll a and SRP concentrations were determined from an 18- meter depth; Emerald Bay chlorophyll a and SRP concentrations were determined from an 11- meter depth. Lake Tahoe chlorophyll a and Emerald Bay secchi measurements were measured in October 2019; all other measurements were performed in July 2019. 1) Schladow 2019………………………………………………………………………………………………..32 2. Description of the 5 treatments in each experiment. Each treatment consisted of 5 replicates. Each mesocosm contained approximately 27 mg dry weight of the study organism…………………………………………………………………………………………...33 3. Results for the Lake Tahoe experiment. The values outside of the parentheses are the percent increase (positive value) or decrease (negative value) in the parameter relative to the Control of the experiment. The values in parentheses are the p-values of the statistical tests. Values marked with a P indicate that a permutation test was used, all other values were determined with a Dunnett’s test. Values that are statistically significant (p < 0.05) are bolded..................................34 4. Results for Emerald Bay experiment. The values outside of the parentheses are the percent increase (positive value) or decrease (negative value) in the parameter relative to the Control of the experiment. The values in parentheses are the p-values of the statistical tests. Values marked with a P indicate that a permutation test was used, all other values were determined with a Dunnett’s test. Values that are statistically significant (p < 0.05) are bolded..................................35 Supplemental Table 1. Relative % error in cumulative particle (diameters ≥ 0.5 µm) concentrations for samples measured in duplicate………………………………………………...43 Chapter 2 1. Limnological variables of the study ecosystems. 1) Dong 1975; 2) Hanes 1981; 3) Morgan 1981..................................................................................................................................................72 Supplemental Table 1. Bulk tissue δ13C and δ15N of mysids……………………………………...80 Supplemental Table 2. Bulk tissue δ15N of herbivorous zooplankton………………………….....84 Supplemental Table 3. Percentages of dietary items in mysid foregut analyses incorporated into the bioenergetics model. Matter classified in the “Other” category consists of unidentifiable matter that was not incorporated into the models…………………………………………………………86 Supplemental Table 4. Amino acid δ15N values of mysids and the calculated trophic positions…89 Supplemental Table 5. Energy density for prey items used in the bioenergetics models…………90 vi List of Figures Chapter 1 1. Chlorophyll a concentrations, pheophytin / chlorophyll a ratios, and biomass-specific PPR. The Lake Tahoe experiment values are shown in the graphs in the left-hand column, and the Emerald Bay experiment values are shown in the graphs in the right-hand column. Treatments that are significantly different (p < 0.05) from the Removal are marked with ** below the x-axis label; treatments that are marginally different (p < 0.1) are marked with *. The mean value of the Control is marked with a dashed line. For each boxplot, the solid horizontal line in the box
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