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«A DISSERTATION SUBMITTED TO THE FACULTY OF THE UNIVERSITY OF MINNESOTA BY Tyler Douglas Ahrenstorff IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR ...»

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Diel Vertical Migrations and Trophic Interactions of Freshwater Organisms

A DISSERTATION

SUBMITTED TO THE FACULTY OF THE

UNIVERSITY OF MINNESOTA

BY

Tyler Douglas Ahrenstorff

IN PARTIAL FULFILLMENT OF THE REQUIREMENTS

FOR THE DEGREE OF

DOCTOR OF PHILOSOPHY

Dr. Thomas R. Hrabik, Advisor May 2013 © Tyler Douglas Ahrenstorff 2013 Acknowledgements I would like to start by thanking my advisor, Tom Hrabik, who successfully guided me through my MS degree and now my PhD. His knowledge, time, patience, and encouragement were instrumental in my accomplishments. I would also like to thank my PhD committee consisting of Donn Branstrator, Stephanie Guildford, and Bob Hecky for all of their advice, guidance, and for borrowing a lot of research equipment to me. In addition, I would like to acknowledge my co-authors, including, Andy Carlson, Beth Holbrook, Pete Jacobson, Olaf Jensen, Dan Yule, Bud Mendsaikhan, Don Pereira, Greg Sass, Jason Stockwell, and Brian Weidel.

I thank the Integrated Biosciences Program and the Biology Department for providing me with teaching assistantships, summer paychecks, and a multitude of other things. It would be impossible for me to list all the faculty and staff that helped me along the way. I appreciate all of the encouragement I have received from fellow graduate students, friends, and other colleagues. Specifically, I would like to thank Josh Dumke, Allison Gamble, EJ Isaac, Tyler Kaspar, Mike Lynch, Mark Mayefske, Samantha Oliver, Kirk Olson, Ryan Oster, and Brian Roth (and probably others I am forgetting). I also want to thank the undergraduates that have helped with research, including, Kyle Gilles, Logan Jacobson, Carl Liebe, Jacob Olson, Thomas Pevan, and Jiethyl Piersak.

Most importantly, I want to thank my wonderful wife, Carla Ahrenstorff, who has given me unending love and support. I also want to acknowledge my parents, Colleen and Doug Ahrenstorff, and my sister, Jessica Graft, who have encouraged me throughout my life.

i Abstract Abiotic and biotic conditions interact to cause dynamic diel vertical migrations (DVM) of organisms in the pelagic area of freshwater lakes. This dissertation includes four unique studies to improve our knowledge of how and why DVM patterns change between species, ecosystems, seasons, and years. The first study documented a normal DVM of the endangered Hovsgol grayling (Thymallus nigrescens) in Lake Hovsgol, Mongolia, during the summer season. This migration was driven by the distribution of prey because there were no pelagic predators. Next, I examined how different abiotic and biotic conditions between 11 inland lakes in Minnesota and northern Wisconsin caused differences in DVM patterns of cisco (Coregonus artedi) during the summer season. I found cisco populations that performed normal DVM, no DVM, and reverse DVM, resulting from differences in temperature, oxygen, prey density, growth potential, and predation risk between lakes. To determine differences in DVM patterns between seasons and years, I followed up this study by analyzing reverse DVM patterns of cisco in Ten Mile Lake, Minnesota, during the spring, summer, fall, and winter over multiple years. I observed a small reverse DVM of cisco during the spring and fall, a large reverse DVM during summer, with no DVM during winter. These differences were attributable to seasonal changes in the abiotic and biotic conditions of the lake. Lastly, I examined seasonal and yearly changes in normal DVM patterns of four pelagic species in Lake Superior. DVM patterns changed between seasons, but were more consistent among years, and were driven by changes in prey and predator densities and distributions.

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Acknowledgements……………………………………………………………………... i Abstract...……………………………………………………………………………….. ii Table of Contents…………………………………………………………………......... iii List of Tables……………………………………………………………………........... iv List of Figures…………………………………………………………………………... v Forward………………………………………………………………………................ x Chapter 1: Introduction………………………………………………..............……….. 1 Chapter 2: Lake Hovsgol, Mongolia……………………………………..............……. 7 Chapter 3: Minnesota and Wisconsin Inland Lakes…………………….............…..... 24 Chapter 4: Ten Mile Lake, Minnesota……………………………………....…........... 44 Chapter 5: Lake Superior, USA……………………………………………….........… 61 Chapter 6: Conclusions……………………………………………………….............. 93 References……………………………………………………………………............. 132

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Table 1. Physical and biological characteristics of the hydroacoustic transects sampled in Lake Hovsgol, Mongolia.





……………………………………………………....... 99 Table 2. Physical and biological attributes of the study lakes sampled in Minnesota and northern Wisconsin. ………………………………………………………....…..… 100 Table 3. Biological and physical parameters used in the foraging, growth, and predation risk models for Lake Superior. ……………………………………………………. 101 Table 4. Weighted, penalized regression spline fits predicting mysis (Mysis diluviana), kiyi (Coregonus kiyi), cisco (C. artedi), and siscowet lake trout (Salvelinus namaycush) depths based on time of day for spring, summer, and fall from 2005 – 2008 in Lake Superior. …………………………………………………………………………… 102

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Figure 1. Areas of Lake Hovsgol, Mongolia, sampled using hydroacoustics, vertical and horizontal gillnetting, and zooplankton net tows.

……………………………....... 103 Figure 2. Length-frequency distributions for Hovsgol grayling sampled with A) horizontal gillnets, B) vertical gillnets, and C) hydroacoustics. ……………......... 104 Figure 3. Diet composition (% by mass) of Hovsgol grayling (Thymallus nigrescens) caught in A) pelagic vertical gillnets and B) littoral horizontal gillnets. ……........ 105 Figure 4. Transect mean density estimates (ind.∙ha-1) of A) smaller (100 – 170 mm) and

B) larger ( 170 mm) Hovsgol grayling (Thymallus nigrescens) in Lake Hovsgol, Mongolia. ……………………………………………………………………….... 106 Figure 5. Day and night vertical distributions of smaller Hovsgol grayling (100 – 170 mm; Thymallus nigrescens) and larger Hovgsol grayling ( 170 mm) at 8 transects (e.g.

T2, T3, etc.) throughout Lake Hovsgol, Mongolia. …………………………….... 107 Figure 6. Day and night vertical migrations, by proportion in each meter of the water column, of the zooplankton community consisting of A) calanoid copepods, B) cyclopoid copepods, C) Daphnia spp., and D) Bosmina spp. ………………………………. 108

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2010-2011 in Minnesota, USA (n=4). ………………………………………........ 109 Figure 8. Length-frequency (mm total length) distributions for cisco captured in each lake during different years. ………………………………………………....…..... 110 Figure 9. Cisco vertical distributions during the day and night in each study lake during different years showing no diel vertical migrations, reverse diel vertical migrations, and normal diel vertical migrations. ………………………………………………...... 111 Figure 10. Taxonomic composition of the diet (proportion by mass) for cisco in each study lake during each year sampled. ……………………………………………. 112 Figure 11. Model performance (% overlap) between the observed and predicted distribution of cisco in Lake Carlos, Minnesota. ……………………………….... 113 Figure 12. Box and whisker plots showing the average percent overlap for the temperature/oxygen, foraging rate potential, growth rate potential, and predation risk models. ………………………………………………………………………….... 114 Figure 13. Model performance (% overlap) between the observed and predicted distributions of cisco in Minnesota and northern Wisconsin lakes. …………..….. 115

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cisco (mm) in each inland lake. ………………………………………………...… 116 Figure 15. Locations of stationary sampling sites and hydroacoustic transects sampled during the summer of 2010 and the spring, summer, fall, and winter of 2011 and 2012 in Ten Mile Lake, Minnesota. ……………………………………………………..... 117 Figure 16. Length-frequency (mm total length) distribution of cisco (Coregonus artedi) in Ten Mile Lake caught using vertical gillnets during the spring of 2011, summer of 2010, 2011, and 2012, and the winter of 2011 and 2012. ………………….…….. 118 Figure 17. Diet composition (proportion by mass) of cisco (Coregonus artedi) during different seasons and years in Ten Mile Lake, MN. …………………………....... 119 Figure 18. Day and night vertical distributions of cisco (Coregonus artedi) in Ten Mile Lake during the spring, summer, fall, and winter of 2010 (dashed line), 2011 (dotted line), and 2012 (solid line). ……………………………………………….……… 120 Figure 19. Foraging (solid line) and predation risk (dotted line) model predictions for cisco (Coregonus artedi) in Ten Mile Lake during the spring, summer, fall, and winter of 2010, 2011, and 2012. ………………………………………………………...….. 121

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fall, and winter of 2010, 2011, and 2012. ………………………………………... 122 Figure 21. Day and night growth model predictions (g/day) of cisco in Ten Mile Lake during the spring, summer, fall, and winter of 2010 (light gray line), 2011 (dark gray line), and 2012 (black line). …………………………………………………….... 123 Figure 22. The survey transect sampled seasonally from 2005-2008 in the western arm of Lake Superior. ……………………………………………………………….... 124 Figure 23. Length-frequency distributions for kiyi (Coregonus kiyi), cisco (C. artedi), and siscowet lake trout (Salvelinus namaycush) captured during all seasons in midwater (from 2003 – 2009) and bottom trawls (from 2007 – 2008). …………………….. 125 Figure 24. Diet proportions calculated by mass for (A) kiyi, (B) cisco, and (C) siscowet lake trout captured in midwater and bottom trawls along the study transect from 2005 – 2008 during each season. …………………………………………...……………. 126 Figure 25. Weighted, penalized regression splines modeling diel vertical migration behavior with time as the predictor, depth the response, and weighted by density during all seasons from 2005 – 2008. ……………………………………………....……. 127

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day and night in three seasons during 2007 – 2008. ……………………………... 128 Figure 27. Seasonal light levels (lux) and water temperature (°C) by depth in Lake Superior. ……………………………………………………………………….…. 129 Figure 28. Kiyi (Coregonus kiyi) observed and predicted depth distributions from the day and night during all seasons in 2007 – 2008. …………………………….….. 130 Figure 29. Cisco (Coregonus artedi) observed and predicted depth distributions from the day and night during all seasons in 2007 – 2008. …………………………….….. 131

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This is a doctoral dissertation submitted as partial fulfillment for the degree of Doctor of Philosophy from the University of Minnesota. Chapter 2 is currently published in Environmental Biology of Fishes. Chapter 3 is currently accepted with revisions in Oecologia. Chapter 5 is currently published in Transactions of the American Fisheries Society. All of the chapters were prepared for publication with co-authors, so the collective “we” is used instead of “I” throughout the dissertation.

Ahrenstorff, T.D., O.P. Jensen, B.C. Weidel, B. Mendsaikhan, and T.R. Hrabik. 2012.

Abundance, spatial distribution, and diet of endangered Hovsgol grayling (Thymallus nigrescens). Environmental Biology of Fishes 94:465-476.

http://link.springer.com/article/10.1007/s10641-011-9961-5#page-1 Ahrenstorff, T.D., T.R. Hrabik, P.C. Jacobson, and D.L. Pereira. Accepted. Food resource effects on diel movements and body size of cisco in north-temperate lakes.

Oecologia. http://www.springer.com/life+sciences/ecology/journal/442 Ahrenstorff, T.D., T.R. Hrabik, J.D. Stockwell, D.L. Yule, and G.G. Sass. 2011.

Seasonally dynamic diel vertical migrations of Mysis diluviana, coregonine fishes, and siscowet lake trout in the pelagia of western Lake Superior.

Transactions of the American Fisheries Society 140:1504-1520.

http://www.tandfonline.com/doi/abs/10.1080/00028487.2011.637004#

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