AQUATIC CONSERVATION BIOLOGY IN ARID ECOSYSTEMS By Eric C. Dinger A Dissertation Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in Biology Northern Arizona University December 2006 Approved: ________________________ Jane Marks, Ph.D., Chair ________________________ Philip Service, Ph.D. ________________________ Tad Theimer, Ph.D. ________________________ Roderic Parnell, Ph.D. ABSTRACT AQUATIC CONSERVATION BIOLOGY IN ARID ECOSYSTEMS Eric C. Dinger Aquatic conservation in arid ecosystems is a pressing concern for the public and land managers in the Southwest. This study focuses on aquatic conservation issues in 2 sites – 1 international site, Cuatro Ciéneags, México and 1 Arizona site, Fossil Creek which was the focus of a collaborative, multi-faceted stream restoration project. In Cuatro Ciénegas, we conducted an experiment manipulating fish access to stromatolites. We manipulated 2 fish species that occur with stromatolites, the polymorphic Herichthys minckleyi, and the pupfish Cyprinidon bifasciatus. We used a trophic cascade index as an indicator of cascade strength, and only molariform morphs were responsible for a trophic cascade, reducing snail densities so that stromatolite algal biomass was positive. The papilliform morph treatments, in contrast, allowed snail densities to increase, resulting in stromatolite algae declines indicating loss of stromatolite formation. Our results show that modern stromatolite formation requires the presence of a specific keystone morph of an endemic threatened cichlid. Our results are also consistent with the hypothesis that metazoan grazing could have been responsible for ancient stromatolite declines, and modern stromatolites should be studied in the context of the entire ecosystem. Restoring native fish to freshwater habitats often requires removal of exotic fish using chemicals such as Antimycin A. We studied the immediate and lingering effects of Antimycin A on macroinvertebrates during a fish renovation project in Fossil Creek, ii Arizona. We employed before-after-control-impact designs to measure the effects of Antimycin A (at 54 µg/L and 100 µg/L) on macroinvertebrate drift, densities and species composition. At the highest dose (100 µg/L) Antimycin A increased drift five fold and immediately decreased invertebrate standing stocks in pools and riffles. Although Antimycin A effects were mostly short-term, several species were extirpated. We studied the short-term effects of restoration of flows to Fossil Creek after 100+ years of flow diversion. Invertebrate density and diversity was unaffected, but there was a rapid response in species composition to flow restoration at restored sites. Downstream sites shifted as a response to flow, but long-term effects will likely be the result of changing geomorphology associated with changing travertine deposition. iii Acknowledgements Dissertation projects do not occur by themselves or through the work of one individual. I received a large amount of help from a wide variety of people – so many I cannot hope to list them all here. First and foremost, thanks to my family (especially my parents) who instilled my love of the natural world at an early age. Thanks also to my high school biology teachers who started me down this path. I am also indebted to the entire graduate community at Northern Arizona University, especially Bill and Anne Stanely, Rusty and Bianca Perla, Matt and Heidi O’Neill, Brook and Lisa Swanson – these people were my role models for my entire graduate career. I can only hope to live to be as good as they are. Thanks to all the people who provided various assistance throughout: Joe Shannon, Bruce Hungate, Alice Gibb, Mike Kearsely, Eric North, Kevin Wilson and Allen Haden. They all provided me with advice or field assistance during my Ph.D. studies, as well as my Masters degree. Special thanks to Dr. Rick Doucett – who help me in so many ways, it’s impossible to remember them all. He single-handedly helped me remember my joy in climbing, running, and beer drinking. And then became a willing partner in all of these pursuits. As a person who disdains paperwork, thanks to the entire Biology office staff who were very accommodating as I dealt with filing various papers! And to Dr. Ron Markle who (usually) was very patient with my questions. My committee was also incredibly patient with me – from my late start in even having a committee to the final rush to finish my degree. Phil Service, Tad Theimer and Rod Parnell were the ideal committee to see me through this. Dr. Jane Marks was my collaborator through all of this – She helped me learn how to be a good scientist, educator, professional and human being. It has been a joy working with Jane from the start of both of our careers – she was the prefect advisor to me. iv Despite this help, I don’t think it would have finished without the help of one last person – my loving wife Nikki. In so many ways, this is not my dissertation, it is our dissertation! Her guidance, support and encouragement provided me the light in the many dark tunnels during my education. v Table of Contents PAGE Dissertation Abstract ii Acknowledgements iv List of Tables viii List of Figures x Dedication xii Preface xiii Chapter 1. Predatory fish maintain modern stromatolites Abstract 1 Text 2 References and Notes 4 Materials and Methods 6 Chapter 2. Modern Stromatolites in their ecosystem: Is modern stromatolite growth maintained by higher trophic levels? Abstract 9 Introduction 10 Methods 12 Results 17 Discussion 19 Literature Cited 24 Chapter 3. Antimycin A affects macroinvertebrates: mortality and recovery Abstract 38 Introduction 40 Study Site 42 Methods 44 Results 50 Discussion 53 vi Table of Contents, continued Literature Cited 58 PAGE Chapter 4. Evidence of rapid responses by macroinvertebrate communities to restored flow in a previously regulated river. Abstract 74 Introduction 75 Methods 78 Results 82 Discussion 83 Literature Cited 88 vii List of Tables TABLE NO. TITLE PAGE Chapter 2. Table 1. Cascade strength results as measured as log ratios. 28 Table 2. Summary of invertebrate means and ANOVA results of dominant orders in Herichthys minckleyi experiment. 28 Table 3. Summary of invertebrate means and ANOVA results of dominant orders in Cyprinodon bifaciatus experiment. 28 Appendix 1. Invertebrates collected from experimental stromatolites during harvest time. 36 Chapter 3. Table 1. Physical and chemical characteristics values for study sites. 61 Table 2. Treatment and sampling schedule for Antimycin A treatment of Fossil Creek 62 Table 3. Results of ANOVA tests at Treatment Site 1 on changes in total invertebrate densities and changes in HBI from Antimycin A treatment. 63 Table 4. Results of ANOVA tests at Treatment Site 2 on changes in total invertebrate densities and changes in HBI from Antimycin A treatment 64 Table 5. Results of indicator species analysis for control and treatment sites showing extirpation of invertebrates following Antimycin A treatment. 65 Chapter 4. Table 1. Sample site characteristics and hypothesized changes for invertebrate assemblages in Fossil Creek, Arizona. 92 Table 2. BACI ANOVA results for density and species richness. 92 Table 3. ANOSIM, SIMPER and BVSTEP results for pre to post-flow restoration changes. 93 viii List of Tables, continued TABLE NO. TITLE PAGE Appendix 1. Taxa list and numbers collected in all 6 sites over the course of the study. 102 ix List of Figures FIGURE NO. TITLE PAGE Chapter 1. Figure 1. Trophic cascade promotes stromatolite growth. 5 Chapter 2. Figure 1. Average invertebrate response to H. minckleyi morph manipulation. 29 Figure 2. Average invertebrate response to H. minckleyi morph manipulation. 30 Figure 3. Average stromatolite algal biomass response to H. minckleyi morph manipulation. 31 Figure 4. Average invertebrate response to C. bifasciatus manipulation. 32 Figure 5. Average invertebrate response to C. bifasciatus manipulation. 33 Figure 6. Average stromatolite algal biomass response to C. bifasciatus manipulation. 34 Figure 7. Stromatolite biomass (as Chl a) for all treatments from both experiments in relation to snail density (Gastropoda). 35 Chapter 3. Figure 1. Map of Fossil Creek, Arizona showing study sites. 66 Figure 2. Mean drift, benthos densities and HBI in Treatment Site 1. 67 Figure 3. Non-Metric Multidimensional Scaling ordination of invertebrate riffle assemblages in Treatment Site 1. 68 Figure 4. Mean drift, benthos densities and HBI in Treatment Site 2. 69 Figure 5. Non-metric Multidimensional Scaling ordination of Treatment Site 2 drift. 70 x