AN ABSTRACT OF THE THESIS OF David B. Herbst for the degree of Doctor of Philosophy in Zoology and Entomology presented on March 6, 1986 . Title: COMPARATIVE STUDIES OF THE POPULATION ECOLOGY AND LIFE HISTORY PATTERNS OF AN ALKALINE SALT LAKE INSECT: EPHYDRA (HYDROPYRUS) HIANS (DIPTERA:EPHYDRIDAE) Redacted for Privacy Abstract approved: -v-- U rank,,P. Conte (Zoology) 1; Redacted for Privacy or J. Brookes (Entomology) Seasonal abundance and life history traits were compared between populations of the brine fly ERhydra (Hydrokyrus) hians Say (Diptera:Ephydridae), from two western Great Basin alkaline salt Lakes. Abert Lake, Oregon, has a relatively low salinity (20-30 g/1 total dissolved solids) and more co-inhabitant benthic species than the higher salinity Mono Lake, California (75-90 g/l). During a period of declining salinities at both lakes, the abundance of this osmoregulating insect decreased at Abert Lake, and increased at Mono Lake. This suggests that abundance may be maximized at intermediate salinities due to biotic limitations imposed by competing and predatory species at dilute salinities, and physiological limitations imposed by osmotic stress at high salinities. Experimental rearing of larvae at high salinities, or reduced algal food supply levels, produced low survival, prolonged development, and smaller size at maturity. When food is not limiting, Mono Lake larvae exhibit greater independence of the inhibitory effects of increased salt concentration compared to larvae from less saline Abert Lake. Selection for enhanced salt tolerance may thus have occurred at Mono Lake, but appears to be limited above 150 g/1 because survival of first instars, and maturation of final instars are impaired at 200 g/l. Besides direct physiological effects, increased salinity also reduces algal growth and may thereby limit food availability to E. hians. Heritable differences in body size exist between populations in addition to environmentally induced changes in growth and size. Abert flies are inherently larger than Mono flies, and develop more rapidly at comparable salinity. Reduction in pupal size severely curtails emergence, and any small-bodied adults that do emerge possess only slight lipid stores, and have a low resistance to starvation. Improvements in algal food supplied to adults increases the proportion of flies reproducing, fecundity, and egg production rates. Reproductive effort has a negative impact on survival only when food is limiting. These results suggest that direct and indirect effects of changing salinity may play an important role in shaping life history patterns and regulating population dynamics of the alkali fly. COMPARATIVE STUDIES OF THE POPULATION ECOLOGY AND LIFE HISTORY PATTERNS OF AN ALKALINE SALT LAKE INSECT: EPHYDRA (HYDROPYRUS) HIANS SAY (DIPTERA:EPHYDRIDAE) by David B. Herbst A THESIS submitted to Oregon State University in partial fulfillment of the requirement for the degree of Doctor of Philosophy Completed March 6, 1986 Commencement June 8, 1986 APPROVED: Redacted for Privacy Prof ess(ory.ogy in Aire(' of major Redac'---t- ed for Privacy ProfesttiAf Ep-Comology in charge of major Redacted for Privacy Chairman of Department c/Zoology Redacted for Privacy Chairman of Department of Entom ogy Redacted for Privacy Dean of Gra ate Schoo< Date thesis is presented March 6, 1986 Typed by David B. Herbst for David B. Herbst ACKNOWLEDGEMENTS Through these many years of study I am grateful to have had the freedom to explore the paths where my interests have led. I owe sincere thanks for guidance and contributions along these paths to my thesis advisors Drs. Frank P. Conte and Victor J. Brookes, and the other members of my committee: Drs. P.B. McEvoy, C.J. Bayne, R.W. Castenholz, C.D. McIntire, N.H. Anderson, and J. Beatty. In addition, other OSU faculty members including Drs. C.E. King, H.K. Phinney, and S.V. Gregory have freely provided their expertise and advice. There are many friends and colleagues without whom my work would have been far less enjoyable and productive, they include: Dan Bean, Dean Blinn, Katie Boula, Tim Bradley, Nick Collins, Gayle Dana, Joe Furnish, John Harris, Eldon Horst, Bob Jellison, Wayne Mathis, Jim McIver, John Melack, Jim Mende, Andy Pollack, Joseph Scheiring, Dave Shuford, Karl Simpson, Stuart Sloan, Scott Stine, Tim Such, Evan Sugden, Peter Vorster, Dave Winkler, and Jeff Witcosky. For their hospitality on my visits to Mono Lake, and their genuine interest and concern I would like to thank David and Sally Gaines, and Meredith and Larry Ford. For their constant loving support, patience, and encouragement I wish to thank all my family Len, Kay, Rebecca, Suzanne, and Dorothy Herbst. I am thankful also to Faye Cummins for sharing all the joys, frustrations, and growth over these years, and for her excellent artwork (Figure II.1). I also wish to acknowledge the funding support provided by the United States Fish and Wildlife Service, Sigma Xi, The American Museum of Natural History, the Oregon State University Department of Zoology, the Mono Lake Foundation, and the Los Angeles Department of Water and Power. Finally, I would like to give tribute to the ancient desert seas of the Great Basin and the allegories of the alkali fly, from which I have gained inspiration and humility on my sojourns. TABLE OF CONTENTS Page I. COMPARATIVE STUDIES OF THE POPULATION ECOLOGY AND LIFE HISTORY PATTERNS OF AN ALKALINE SALT LAKE INSECT: EPHYDRA (HYDROPYRUS) HIANS SAY (DIPTERA:EPHYDRIDAE) INTRODUCTION 1 II. AUTECOLOGY AND NATURAL HISTORY OF AN ALKALINE SALT LAKE INSECT: EPHYDRA (HYDROPYRUS) HIANS SAY (DIPTERA :EPHYDRIDAE) 6 Introduction 6 Methods 9 Benthic Sampling 9 Temperature and Salinity Effects on Development 10 Results and Discussion 13 Life Cycle and Natural History of the Alkali Fly 13 Microhabitat Associations at Mono Lake 19 Temperature and Salinity Effects 20 III. BENTHIC SPECIES DIVERSITY AND PHENOLOGY IN RELATION TO THE PHYSICAL AND CHEMICAL ENVIRONMENTS OF GREAT BASIN ALKALINE SALT LAKES 31 Introduction 31 History and Physicochemical Environments of Abert and Mono Lakes 32 Methods 37 Sample Sites 37 Temperature and Salinity 38 Benthic Fauna 39 Benthic Flora 40 Results 42 Salinity 42 Temperature 43 Benthic Fauna 43 Benthic Flora 46 Discussion 49 IV. COMPARATIVE POPULATION ECOLOGY OF EPHYDRA HIANS SAY (DIPTERA:EPHYDRIDAE) IN ALKALINE SALT LAKES OF THE WESTERN GREAT BASIN 65 Introduction 65 Methods 68 Relative Abundance Censusing 68 Body Size 70 Reproductive Condition 70 Results 72 Population Dynamics of Ephydra hians ... 72 Body Size 74 Reproductive Condition 74 Discussion 76 V. DEVELOPMENTAL EFFECTS OF SALINITY AND ALGAL FOOD AVAILABILITY ON THE ALKALI FLY EPHYDRA HIANS (DIPTERA:EPHYDRIDAE) 98 Introduction 98 Methods 101 Population Sources 101 Adult Rearing and Egg Collection 101 Osmotic Stress on Larval Development 102 Food Supply Restrictions on Larval Development 105 Results 107 Effect of Osmotic Stress on Larval Development 107 Effect of Food Restriction on Larval Development and Survivorship 109 Influence of Pupal Size on Adult Emergence 109 Discussion 111 VI. THE INFLUENCE OF FOOD AND BODY SIZE ON REPRODUCTION AND LONGEVITY OF AN ALKALINE LAKE SHORE FLY, EPHYDRA HIANS SAY (DIPTERA:EPHYDRIDAE) 125 Introduction 125 Methods 129 Field Collection 129 Laboratory Rearing 130 Experimental Design 131 Eggs and Lipid Content in Flies from Field Populations 134 Statistical Analysis 135 Results 136 Starvation Effects 136 Male Longevity 136 Female Longevity 137 Fecundity and Stored Lipids of Field-Collected Flies 139 Reproduction in Experimental Feeding Studies 140 Timing of Reproduction 142 Correlation Among Adult Life History Traits 144 Discussion 146 VII. GROWTH OF THE FILAMENTOUS GREEN ALGA CTENOCLADUS CIRCINNATUS BORZI (CHLOROPHYCEAE) IN RELATION TO ENVIRONMENTAL SALINITY 171 Introduction 171 Methods 174 Algae Collection and Cloning 174 Culture Media 174 Experimental Inoculations 175 Culture Harvesting 176 Chlorophyll and Water Content 177 Results 178 Effect of Salinity on Growth 178 Chlorophyll Content 179 Cell Form and Water Content 180 Discussion 183 BIBLIOGRAPHY 196 APPENDICES Appendix 1. Sample site habitat descriptions.. 202 Appendix 2. Mono Lake: Seasonality and composition indices of benthic algae 204 Appendix 3. Abert Lake: Seasonality and composition indices of benthic algae 205 Appendix 4. Biochemical composition of Ephydra hians 206
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