Ascendency As an Ecological Indicator in the Nitrogen Flow
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ASCENDENCY AS AN ECOLOGICAL INDICATOR IN THE NITROGEN FLOW OF THE NEUSE RIVER ESTUARY, NORTH CAROLINA, USA by MARC E. NOVAK (Under the Direction of John R. Schramski) ABSTRACT Ascendency, a holistic measure of a system’s aggregate activity and flow distribution, is employed to evaluate sixteen seasonal models of nitrogen flow in the Neuse River Estuary. The results and methods are contrasted with ascendency’s typical use with carbon food web models. Ascendency (A), developmental capacity (C), A/C ratio, total system throughput, average mutual information, overheads, and Finn’s cycling index values for the Neuse are reported. The use of the A/C ratio, rather than ascendency alone, for ecosystem comparisons is supported as found in other recent studies. An indicator ratio (Ag+d) is presented as an alternative to ascendency for short-term analysis, distinguishing between increases in ascendency via growth (g) and development (d), and increases coupled with information loss. Focusing on ascendency as an ecological indicator, an attempt is made to use ascendency theory to inform ecological managers while employing the Neuse River Estuary’s nitrogen flow to inform ascendency theory. INDEX WORDS: Ascendency, Ecological Indicator, Environmental Management, Eutrophication, Goal Function, Modeling, Network Analysis, Nitrogen Cycling. ASCENDENCY AS AN ECOLOGICAL INDICATOR IN THE NITROGEN FLOW OF THE NEUSE RIVER ESTUARY, NORTH CAROLINA, USA by MARC E. NOVAK B.S., The University of Georgia, 1999 A Thesis Submitted to the Graduate Faculty of The University of Georgia in Partial Fulfillment of the Requirements for the Degree MASTER OF SCIENCE ATHENS, GEORGIA 2011 © 2011 Marc E. Novak All Rights Reserved ASCENDENCY AS AN ECOLOGICAL INDICATOR IN THE NITROGEN FLOW OF THE NEUSE RIVER ESTUARY, NORTH CAROLINA, USA by MARC E. NOVAK Major Professor: John R. Schramski Committee: Bernard C. Patten E. William Tollner Electronic Version Approved: Maureen Grasso Dean of the Graduate School The University of Georgia December 2011 iv DEDICATION To Clayton C. Smith, the only person I have encountered who believed unconditionally that I could do no wrong. In this most egregious of errors, teaching me that sometimes trust precedes being trustworthy. v ACKNOWLEDGEMENTS This thesis would not have been completed without the constant support and level-headed encouragement of my major professor, Dr. John R. Schramski. I walked into his office with doubts and walked out with a plan, and I have not looked back since. I would like to thank my committee members, Dr. Bernard C. Patten and Dr. E. William Tollner for reviewing my work and serving on my committee despite overwhelmingly busy schedules. I would also like to thank Dr. David K. Gattie for encouraging me to pursue a Master’s Degree, granting me access to the Gattie Lending Library, and welcoming me to the engineering family. Special thanks to the staff at the University of Georgia’s Department of Engineering, in particular Sherry Wrona who has made the impossible possible and saved me on countless occasions. I would like to thank Rebecca Wiessner and all my friends for their support and for putting up with my ecosystem ramblings. Finally, I thank my parents, Ed and Lora Novak whose financial contributions, counseling, and unwavering support made this possible. vi TABLE OF CONTENTS ACKNOWLEDGEMENTS .............................................................................................................v GLOSSARY OF MATHEMATICAL SYMBOLS ..................................................................... viii LIST OF TABLES ......................................................................................................................... ix LIST OF FIGURES .........................................................................................................................x CHAPTER 1 INTRODUCTION ...................................................................................................1 PREFACE ....................................................................................................1 PURPOSE OF STUDY................................................................................1 NEUSE RIVER ESTUARY ........................................................................3 NITROGEN CYCLING ..............................................................................4 NEUSE RIVER ESTUARY MODEL .........................................................5 FIGURES .....................................................................................................7 2 LITERATURE REVIEW ......................................................................................10 NETWORK ANALYSIS ...........................................................................10 ASCENDENCY THEORY .......................................................................13 NEUSE RIVER ESTUARY RESEARCH ................................................14 3 METHODOLOGY ................................................................................................18 CALCULATING ASCENDENCY ...........................................................18 CHANGES IN ASCENDENCY................................................................20 NETWRK 4.2 ............................................................................................21 FIGURES ...................................................................................................23 vii 4 RESULTS AND DISCUSSION ............................................................................26 THROUGHPUT, INFORMATION AND ASCENDENCY .....................26 DEVELOPMENTAL CAPACITY AND CONSTITUENTS ...................27 CONNECTANCE AND CYCLING .........................................................27 TEMPORAL COMPARISONS ................................................................28 SPATIAL COMPARISONS ......................................................................29 MODEL CONSIDERATIONS ..................................................................29 INTERPRETING CHANGES IN ASCENDENCY ..................................31 FIGURES ...................................................................................................33 TABLES ....................................................................................................40 5 CONCLUSIONS AND FUTURE DIRECTIONS .................................................41 OBJECTIVES ............................................................................................41 FUTURE DIRECTIONS ...........................................................................42 CONCLUSIONS........................................................................................43 REFERENCES ..............................................................................................................................44 APPENDIX A NEUSE RIVER ESTUARY’S NITROGEN FLOW DATA .................................49 B NEUSE RIVER ESTUARY’S ASCENDENCY, TOTAL SYSTEM THROUGHPUT, AVERAGE MUTUAL INFORMATION, AND A/C RATIO ............................................................................................................58 C NEUSE RIVER ESTUARY’S SUPPLEMENTARY INDICES ...........................59 viii GLOSSARY OF MATHEMATICAL SYMBOLS Symbol Definition First Use FCI Finn’s Cycling Index ................................................................................................4 NEA Network Environ Analysis .................................................................................... 11 I/D The I/D ratio (Indirect Effects / Direct Effects) ....................................................12 A/C The A/C ratio (Ascendency / Developmental Capacity) .......................................13 AMI Average Mutual Information .................................................................................18 TST Total System Throughput .......................................................................................18 T.. Flow from any arbitrary compartment to any other ..............................................19 T.j Flow from any arbitrary compartment to compartment j ......................................19 Ti. Flow from compartment i to any arbitrary compartment ......................................19 Tij Flow from compartment i to compartment j .........................................................19 Ti All flows leaving compartment i ...........................................................................19 T'i All flows entering compartment i .........................................................................19 Tik Flow from compartment i to compartment k ........................................................19 Ei Exports from compartment i .................................................................................19 Di Imports to compartment i ......................................................................................19 Dj Imports to compartment j ......................................................................................19 Ri Dissipation from compartment i ...........................................................................19 Ag+d Growth and Development Ratio ...........................................................................32 S1 Number of observations of scenario #1 .................................................................32 Stotal Total number of observations ...............................................................................32 ix LIST OF TABLES Table 4.1: Ascendency and A/C values in geographically