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ANTHROPOGENIC AND LANDSCAPE FACTORS CONTROL STREAM NITROGEN TRANSFORMATIONS AT MULTIPLE SPATIAL SCALES A Dissertation Submitted to the Graduate School of the University of Notre Dame in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy by Clay P. Arango, B.A., M.S. ________________________________ Jennifer L. Tank, Director Graduate Program in Biological Sciences Notre Dame, Indiana July 2007 ANTHROPOGENIC AND LANDSCAPE FACTORS CONTROL STREAM NITROGEN TRANSFORMATIONS AT MULTIPLE SPATIAL SCALES Abstract by Clay P. Arango Anthropogenic alterations to the global nitrogen (N) cycle have doubled reactive N flux into the biosphere and altered aquatic ecosystem function. Streams modify N loads carried to coastal ecosystems by converting N to organic forms or removing it as gaseous N. Understanding how streams transform N can offer insight to ecologists and land managers about how stream ecosystems function under elevated N loads. I researched how anthropogenic and landscape factors affect N transformations by studying streams in two distinct biomes and at multiple spatial scales. At the landscape scale, I studied N concentrations in streams draining the Teton Range (Wyoming, USA), a sub-alpine and alpine ecosystem with variable lithology. Streams draining crystalline geology had higher N compared to streams draining carbonate geology, which had more vegetation, suggesting that lithology mediated patterns in vegetation and terrestrial N retention. At the reach scale, I studied how land use influenced N uptake and transformation in Midwestern streams (Michigan, USA) and found that dissimilatory N transformation rates (i.e., nitrification and denitrification) Clay P. Arango within streams were not affected by riparian zones, which are commonly used to mitigate water quality degradation. Dissimilatory N transformation rates were always < 10% of whole-stream N uptake and nitrification rates balanced denitrification rates, implying that denitrification did not represent net N loss from the water column. At the substratum scale, sediment organic carbon content correlated with denitrification, but only when nitrate concentration exceeded a threshold. Finally, I returned to the Tetons and found that grazing activity by invasive snails can increase periphyton N fixation rates in a stream with low N concentrations. A synthesis of my findings from high N streams in the Midwest suggests that land-use practices have increased temporary N removal at the expense of permanent N removal. In the low N streams of the Tetons, observations from different spatial scales suggest that landscape factors that lower stream N concentrations and high rates of grazing together can influence the importance of N fixation in streams. My dissertation highlights multiple constraints on N processing in streams and emphasizes that basic ecological research can yield important results for management agencies. CONTENTS FIGURES................................................................................................................................. vi TABLES .................................................................................................................................. xi ACKNOWLEDGMENTS ..................................................................................................... xiii CHAPTER 1 DISSERTATION INTRODUCTION .................................................................1 1.1 Human Activity and Global Change.......................................................................1 1.2 Nitrogen Biogeochemistry: Assimilatory and Dissimilatory Processes................2 1.3 Anthropogenic Alterations to the N Cycle .............................................................5 1.4 Nitrogen Cycle in Streams......................................................................................6 1.5 Anthropogenic Alterations to Stream Ecosystems .................................................8 1.6 A Conceptual Model of N Dynamics Incorporating Multiple Spatial Scales ........9 1.7 Dissertation Outline..............................................................................................12 CHAPTER 2 LANDSCAPE CONTROLS ON NITROGEN CONCENTRATIONS IN STREAMS DRAINING THE TETON RANGE, WYOMING ..............................................16 2.1 Abstract.................................................................................................................16 2.2 Introduction...........................................................................................................17 2.3 Methods ................................................................................................................20 2.3.1 Study Site.............................................................................................20 2.3.2 Sample Collection................................................................................20 2.3.3 Water Chemistry Analyses ..................................................................22 2.3.4 GIS Analyses.......................................................................................23 2.3.5 Statistical Analyses ..............................................................................24 2.4 Results...................................................................................................................25 2.4.1 Landscape Factors................................................................................25 2.4.2 Solute Concentrations..........................................................................28 2.4.3 Relationship between Solute Concentrations and Topography, Land Cover, and Lithology..................................................................33 2.4.4 Within-basin Solute Patterns................................................................35 2.4.5 Indexes of Sensitivity to N Saturation .................................................39 2.5 Discussion.............................................................................................................40 2.5.1 Landscape Factors and N Concentrations............................................40 2.5.2 Landscape Factors and Dissolved Organic Matter ..............................43 ii 2.5.3 Ecosystem N Status..............................................................................45 2.6 Acknowledgements...............................................................................................47 CHAPTER 3 LAND USE INFLUENCES THE SPATIO-TEMPORAL CONTROLS ON NITRIFICATION AND DENITRIFICATION IN HEADWATER STREAMS....................48 3.1 Abstract.................................................................................................................48 3.2 Introduction...........................................................................................................49 3.3 Methods ................................................................................................................53 3.3.1 Land-use Classification........................................................................53 3.3.2 Sediment Nitrification Assays .............................................................57 3.3.3 Sediment Denitrification Assays..........................................................58 3.3.4 Sediment Characterization...................................................................60 3.3.5 Physiochemical Variables and Water Chemistry.................................61 3.3.6 Statistical Analyses ..............................................................................61 3.4 Results...................................................................................................................62 3.4.1 Land-use and Seasonal Patterns in N Transformations .......................62 3.4.2 Land-use and Seasonal Patterns of DIN and Organic C......................66 3.4.3 Factors Controlling N Transformation Rates.......................................69 3.4.4 Relationship between Continuous Land-use Metrics and N Transformations ...................................................................................73 3.5 Discussion.............................................................................................................74 3.5.1 Seasonal Patterns of Sediment Nitrification Rates ..............................74 3.5.2 Land-use Influences on Nitrification Controls ....................................76 3.5.3 Seasonal Patterns of Sediment Denitrification Rates...........................78 3.5.4 Land-use Influences on Denitrification Controls.................................79 3.5.5 Effect of Riparian Buffers on Nitrification and Denitrification...........81 3.6 Acknowledgements...............................................................................................85 CHAPTER 4 ASSIMILATORY UPTAKE RATHER THAN DISSIMILATORY TRANSFORMATIONS INFLUENCE SEASONAL PATTERNS IN NITROGEN UPTAKE IN STREAMS OF VARYING LAND USE...........................................................86 4.1 Abstract.................................................................................................................86 4.2 Introduction...........................................................................................................87 4.3 Methods ................................................................................................................91 4.3.1 Land-use Classification........................................................................91 4.3.2 Nutrient Releases.................................................................................92