The Maintenance and Generation of Freshwater Diversity from the Local to the Global Scale
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THE MAINTENANCE AND GENERATION OF FRESHWATER DIVERSITY FROM THE LOCAL TO THE GLOBAL SCALE By Patrick J. Hanly A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of Integrative Biology – Doctor of Philosophy Ecology, Evolutionary Biology, and Behavior – Dual Major 2017 ABSTRACT THE MAINTENANCE AND GENERATION OF FRESHWATER DIVERSITY FROM THE LOCAL TO THE GLOBAL SCALE By Patrick J. Hanly The distribution of biological diversity is markedly uneven across the world. Despite the seemingly endless variety of forms and adaptations that have evolved and continue to evolve, large differences in the numbers and characteristics of species remain among locales. These differences are often not random; e.g., the Earth’s tropics are disproportionately rich in the diversity of life and large areas harbor more species than small areas. Observations of these general biogeographic patterns are some of the oldest contributions by early naturalists and ecologists, yet explanations for these patterns are a recurring topic of debate. Generality of pattern (with room for exceptions to the rule) has been reached for large-scale gradients in diversity but not generality of the theories that underlie these patterns. Unlike diversity at the biogeographic scale, even the generality of pattern at the local scale of species interactions remains elusive. Although numerous investigations into how potential drivers of local diversity such as productivity, isolation, and disturbance influence diversity have been made, a unifying consensus to describe general patterns of local diversity has not emerged. Today, increasing emphasis is being placed on the importance of the interaction between local and regional scales in influencing local diversity. The thesis first introduces and summarizes attempts to describe and explain biodiversity patterns at both the local and regional scales. Chapter 1 describes a study of the role of dispersal and assembly history in influencing species diversity in natural and experimental pond communities of plankton under the metacommunity framework. A further test of the role of dispersal and assembly history is presented in Chapter 2 using experimental pond mesocosms. This study evaluates the role of ecosystem size on the assembly and structure of a multitrophic community of both plankton and macroinvertebrates while concurrently varying nutrient input rate and initial assembly. Chapter 3 empirically illustrates variation in the dormancy-dispersal strategies used by freshwater zooplankton that can lead to interspecific differences in the degree and type of dispersal limitation. In Chapter 4, I use the distribution of single lake endemic fish in the largest lakes in the world to estimate the relative contributions of lake surface area, age, and latitude on diversification. I dedicate this dissertation to my parents, Joseph and Ernestine. iv ACKNOWLEDGMENTS I am grateful for the years of support, guidance, and discussion from my advisor, Gary Mittelbach. May I be as giving to others, as warm-hearted, and as dedicated to the scientific pursuit in my own career. I also acknowledge comments and critique from my other committee members, Elena Litchman, Christopher Klausmeier, and Orlando Sarnelle. Finally, I thank my wife, Melanie, for her selfless support of me in all things. v TABLE OF CONTENTS LIST OF TABLES viii LIST OF FIGURES x INTRODUCTION 1 Chapters 1 & 3 1 Chapter 2 2 Chapter 4 4 LITERATURE CITED 6 CHAPTER ONE 10 THE INFLUENCE OF DISPERSAL ON THE REALIZED TRAJECTORY OF A POND METACOMMUNITY 10 Abstract 10 Introduction 11 Methods 14 Experimental design 14 Statistical analyses 17 Results 22 Natural pond community trajectory 22 Density dependence of plankton dispersal 23 Experimental mesocosm assembly 24 Discussion 25 Acknowledgements 33 APPENDIX 34 LITERATURE CITED 46 CHAPTER TWO 52 THE INFLUENCE OF ECOSYSTEM SIZE ON TROPHIC STRUCTURE AND DIVERSITY DURING AQUATIC COMMUNITY ASSEMBLY 52 Abstract 52 Introduction 54 Methods 56 Mesocosm experiment 56 Data formatting and manipulation prior to analysis 59 Structural equation modeling 60 Endpoint comparison of composition 60 Results 61 Effect of ecosystem size, nutrient input, and assembly status on diversity 61 Influences on food web structure 62 Temporal trends of effects on diversity and trophic structure 63 The observed species-area relationship 65 vi Discussion 65 Effect of ecosystem size on species richness 65 Effect of ecosystem size on trophic structure 69 Lack of nutrient input level effect 70 Conclusion 71 APPENDIX 73 LITERATURE CITED 99 CHAPTER THREE 104 A NEW LOOK AT THE ROLE OF EPHIPPIA AS A DORMANCY-DISPERSAL STRATEGY IN THE CLADOCERAN (SCAPHOLEBERIS MUCRONATA) 104 Abstract 104 Introduction 106 Methods 109 Survey of natural ponds and cladoceran dispersal assessment 109 Outdoor mesocosm experiment 109 Ephippia collection and hatching 112 Statistical analyses 112 Results 113 Pond survey 113 Mesocosm experiment 114 Discussion 116 APPENDIX 120 LITERATURE CITED 126 CHAPTER FOUR 130 SPECIATION AND THE LATITUDINAL DIVERSITY GRADIENT: INSIGHTS FROM THE GLOBAL DISTRIBUTION OF ENDEMIC FISH 130 Abstract 130 Introduction 131 Methods 134 Database construction 134 Analyses on lakes with age data 136 Multiple missing data imputation on the full lake dataset 136 Comparison to fish endemism patterns in rivers 138 Phylogenetic considerations and potential sampling bias 138 Results 140 Discussion 142 Acknowledgements 147 APPENDIX 148 LITERATURE CITED 171 vii LIST OF TABLES Table 1. Permutational multivariate analysis of variance (PERMANOVA) output table evaluating whether plankton composition differed between local communities and dispersers, as well whether the composition of dispersers was significantly different across sampling periods and location in the metacommunity. P-values < 0.05 are in bold. 35 Table 2. Similarity Percentage (SIMPER) results showing the species responsible for the greatest difference between the composition of local communities and dispersers, along with their percent contribution to this difference. The top ten species underrepresented in the dispersers relative to their metacommunity occupancies are shown above and the top ten overrepresented species are shown below. 36 Table 3. Fitted models describing the relationship between the metacommunity abundance of taxa within eight plankton taxonomic groups and their measured dispersal rates throughout the growing season. P-values < 0.05 are in bold. 37 Table 4. Linear model of the influence of treatments on the final macroinvertebrate species richness of communities. 74 Table 5. Linear model of the influence of treatments on the final macroinvertebrate density of communities. 75 Table 6. Linear model of the influence of treatments on the final zooplankton species richness of communities. 76 Table 7. Linear model of the influence of treatments on the final zooplankton density of communities. 77 Table 8. PERMANOVA of final zooplankton communities. 78 Table 9. PERMANOVA of final macroinvertebrate communities in the two largest mesocosms sizes. 79 Table 10. Slopes of the species-area relationship derived from the literature for freshwater groups with the range of surface areas used in each study. NA indicates a lack of a species-area relationship. 93 Table 11. List of taxa observed in the mesocosm experiment. 98 Table 12. Generalized linear model of the probability that a lake contains an endemic based on a missing data imputation of predictor variables for all 2,746 lakes. 160 viii Table 13. Generalized linear model of the number of endemics in a lake based on a missing data imputation of predictor variables for all 2,746 lakes. 161 Table 14. Generalized linear model of the probability that a river basin contains an endemic fish. Data from the Fish-SPRICH database. 162 Table 15. Generalized linear model predicting the number of endemic fish in a river basin. Data from the Fish-SPRICH database. 163 Table 16. Generalized linear model of the probability of a lake containing an endemic fish with the three largest African Great Lakes removed. 164 Table 17. Generalized linear model of the number of endemic fish in lakes with endemics with the three largest African Great Lakes removed. 165 Table 18. Generalized linear model of the probability of a lake containing an endemic fish with the eight largest African Great Lakes removed. 166 Table 19. Generalized linear model of the number of endemic fish in lakes with endemics with the eight largest African Great Lakes removed. 167 Table 20. Generalized linear model of the probability of a lake containing an endemic fish with the family Cichlidae removed. 168 Table 21. Generalized linear model of the number of endemic fish in lakes with endemics with the family Cichlidae removed. 169 Table 22. Distributional centers of the non-endemic native species and endemic fish for the 47 families containing at least one endemic fish species or subspecies in the 2,746 largest lakes in the world. 170 ix LIST OF FIGURES Figure 1. Non-metric multidimensional scaling (NMDS) ordination plot showing the difference in composition between local communities (green circles) and dispersers (blue crosses) across all sampling periods. The composition of the ten local communities on the final sampling date (i.e., the final metacommunity) is represented by the filled circles (black), which show a visible attraction toward the composition of dispersers that is confirmed by the subsequent analysis using the attraction coefficient (AC). 38 Figure 2. Boxplots