<<

EFFECTS OF A PULSING HYDROPERIOD ON A CREATED RIPARIAN DIVERSION WETLAND

DISSERTATION

Presented in Partial Fulfillment of the Requirements for

the Degree Doctor of Philosophy in the Graduate

School of the Ohio State University

By

Daniel F. Fink, M.S.

* * * * *

The Ohio State University

2007

Dissertation Committee: Approved by

William J. Mitsch, adviser

M. Siobhan Fennessy ______Jay F. Martin Adviser

Franklin W. Schwartz Environmental Science Graduate Program

ABSTRACT

In this study, succession, hydroperiod, and water-quality dynamics are documented for a whole-ecosystem study involving a 3-ha created riparian wetland at the

Schiermeier Olentangy River Wetland Research Park at The Ohio State University in

Columbus, Ohio USA during 2003 through 2006. The effect of delivering influent water to the wetland as seasonal pulses or as a continuous steady flow on the hydroperiod, water chemistry, avian use, plant primary production, and plant community structure within the wetland was investigated. A simulation model was then developed using short-term measurements of river and wetland stage to predict long-term patterns of succession and plant development within the wetland. This wetland typically receives seven or eight natural pulses each year from the Olentangy River. Of 21 species planted in 1997, only Scirpus americanus and Juncus effusus remained as important macrophyte species during the study six to seven years after planting. Typha spp.

(angustifolia and latifolia), colonized naturally and was the dominant macrophyte in most of the wetland during this study.

Mean nutrient reductions per flood pulse for nitrate-nitrite, total Kjeldahl nitrogen

(TKN), soluble reactive phosphorus (SRP), and total phosphorus (TP) during 2003-2004 were 0.71 g-N m-2, 0.92 g-N m-2, 0.016 g-P m-2, and 0.08 g-P m-2 respectively. The

- annual reductions of N-NO3 , TN, P-SRP, and TP were 74%, 41%, 46%, and 31% by

ii - mass. A greater removal of NO3 and TP occurred in the emergent marsh section of the wetland than the open water section. Conversely TKN increased through the emergent marsh and decreased through the open water. Overall, the oxbow design appeared to be successful in ecological terms and similar diversion wetlands are recommended for other locations to examine their function under different climates and hydroperiods.

While hydrologic pulses affect wetland function and may enhance productivity and contribute stability to ecosystems, the overall effects of these pulses on biogeo- chemical processes in riparian river diversion wetland ecosystems have not been clearly demonstrated experimentally at an ecosystem scale. The natural hydroperiod of the riparian wetland was varied in 2005; its natural flood pulses were removed and replaced with an artificial steady-flow supplied by submersed pumps. The wetland received 8 natural river pulses from April 2004 – March 2005, then a steady flow of river water was pumped through the wetland from April 2005 – March 2006. In 2004 the wetland received a total of 27 m3 m-2 yr-1 (cubic meters of influent water per square meter of wetland area per year) of inflow followed by 20 m3 m-2 yr-1 of inflow in the steady flow year 2005. The removal rate was higher for nitrate-nitrogen, total nitrogen, and total phosphorus during the year with flood-pulsing than during the steady-flow year. The only nutrient that did not have a difference in removal rate in the two years was soluble reactive phosphorus. There were differences in spatial dynamics of most of the nutrients in flood-pulsing and non-pulsing years and between wet and dry seasons in all years of the study. In all cases, however, total nitrogen concentrations in the wetland increased through the emergent marsh and then decreased across the open water basin.

iii There was greater avian use of the created oxbow wetland during pulsing than in steady flow conditions. The guild of bird species most affected by the removal of a pulsing hydroperiod was shorebirds, which had a 75% reduction in the number of species observed during steady-flow. Peak observed shorebird use during the pulsing hydroperiod year corresponded to dry conditions in the wetland in the later part of the growing season during shorebird migration time.

Primary productivity of macrophytes was compared among the 2004 pulsing hydroperiod year, the 2005 stead-flow year, and in 2006 when natural pulsing was again restored (6 natural river pulses occurred from April 2006 – September 2006). The wetland was significantly more productive (α = 0.05) during pulsing compared to steady flow conditions. Wetland macrophyte productivity decreased from 7,560 kg yr-1`to 6,544 kg yr-1and total macrophyte areal coverage decreased from 32.3% to 25.7% of the wetland after the pulsing hydroperiod was replaced by the steady-flow hydroperiod.

When a flood pulse hydroperiod was restored in 2006, total net macrophyte productivity rebounded to 9,916 kg yr-1 and areal coverage returned to 32.1%. There was a positive correlation between the flashiness of the wetland hydroperiod and productivity (r = 0.75).

A pulsing hydroperiod should enable mitigation wetlands to reach target macrophyte coverage goals more easily.

A simulation model was developed use short-term hydrologic data, such as the data collected in this study, to predict long-term successional development in a created riparian wetland ecosystem. A hydrologic submodel predicted wetland water stage and depth based on precipitation, potential evapotranspiration, river stage, overland outflows, and groundwater exchange. Vegetation submodels calculated the growth of trees and

iv emergent macrophytes based on water depth and the duration and timing of flooding.

The model also predicted the state of the ecosystem annually as being open water, mudflat, marsh, or forested wetland. Data from 2004 were used to calibrate the models.

Simulation results were compared with actual data from 2003, 2005, and 2006 to validate the model. When the wetland inflow was pulsing, the model predicted water depth within 6.0 cm 75% of the time and when it was a steady flow the model predicted water depth within 2.5 cm 75% of the time.

Simulations of floral succession were evaluated at 8 ( wetland age) and

100 years. Simulations for current wetland age correctly predicted the state of the ecosystem. One hundred year simulations showed annual variations in macrophyte coverage over much of the wetland, but a consistent growth of trees on the wetland fringe. Simulations also predicted a larger area of mudflat if flashiness increased and a greater area of open water if flashiness decreased or if beavers dammed the outflow.

When a accretion routine was added to the model, 100-yr simulations predicted that the wetland will eventually become completely covered with macrophytes and that the area of forested wetland will cover not only the outer fringe, but much of the internal portion of the wetland basin as well. Adding several model features including a dispersal model for propagules and a shorter time step for vegetation could improve model function.

v

Dedicated to my family and friends, who have cared so much for me.

vi ACKNOWLEDGEMENTS

I gratefully thank my advisor, William Mitsch for his wisdom and his patience while guiding me through this process. I also thank Siobhan Fennessy, Jay Martin, and

Frank Schwartz for serving on my dissertation committee and for their support and insight. The quality of this project is much better for their contributions. I am also grateful to my fellow “wetlanders” throughout my stay at Ohio State. Their help and friendship made my time here at The Ohio State University a joyful time in my life.

Funding for my education and research came from The Environmental Sciences Graduate

Program, the National Science Foundation’s GK-12 Fellowship program, the U.S.

Department of Agriculture NRI CSREES Award 2003-35102-13518 and a Payne grant from the Ohio Agricultural Research and Development Center of The Ohio State

University. Lastly I thank my friends and family for supporting me through my long journey in academia. I especially thank my wife Colleen for her love, support, and patience. I have never been happier then since you have been in my life.

vii VITA

12 December, 1975………………… Born – Elyria, Ohio

1998………………………………... B.A., Physics and Environmental Studies

Ohio Wesleyan University

Delaware, Ohio

1998-2001………………………….. Graduate Teaching Assistant,

The Ohio State University

2001………………………………... M.S. Environmental Science Graduate

Program, The Ohio State University

Columbus, Ohio

2002 – 2003………………………... Graduate Teaching and Research Assistant

The Ohio State University, Columbus Ohio

2003 – 2004………………………... GK-12 National Science Foundation

Teaching Fellow, Columbus, Ohio

2005 – 2006………………………... Graduate Teaching and Research Assistant

The Ohio State University, Columbus Ohio

viii PUBLICATIONS

Peer-reviewed journal articles

Fink, D.F. and W.J. Mitsch. 2004. Seasonal and storm event nutrient removal by a created wetland in an agricultural watershed. Ecological Engineering 23:313-325.

Fink, D.F. and W.J. Mitsch. in press. Hydrology and nutrient biogeochemistry in a created river diversion oxbow wetland. Ecological Engineering.

Zhang, L., W.J. Mitsch, and D.F. Fink. 2004. Hydrology, water quality, and restoration potential for the Upper Big Darby Creek, Central Ohio. Ohio Journal of Science 105:46-56.

Publications in the Olentangy River Wetland Research Park Annual Reports

Anderson, C.J., D.F. Fink, W.J. Mitsch. 2003. Vegetation establishment in the mitigation billabong at the Olentangy River Wetland Research park, 2000-2002. In: Mitsch, W.J., L. Zhang, C.J. Anderson (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2002, pp. 101-106

Gilbert, J.M., D.F. Fink, and M. Greene. 1999. Soil properties of three newly created wetlands. In: Mitsch, W.J., V. Bouchard (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 1998, pp. 113-118.

Fink, D.F. and W.J. Mitsch. 2000. Effectiveness of a newly constructed wetland on agricultural run-off. In: Mitsch, W.J., L. Zhang (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2001, pp. 191-196.

Fink, D.F. and W.J. Mitsch. 2001. Wetlands for controlling nonpoint source from agriculture: Indian Lake Wetland Demonstration Project, Logan County, OH. In: Mitsch, W.J., L. Zhang (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2000, pp. 161-178.

Fink, D.F. and W.J. Mitsch. 2005. Fish and amphibian abundance in created riparian marshes with pulsing hydrology. In: Mitsch, W.J., L. Zhang, and A.E. Altor (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2004, pp. 125-128.

Fink. D.F. and W.J. Mitsch. 2005. Hydrology, biogeochemistry, and plant community development in a created river diversion oxbow wetland in the Ohio River Basin, USA. In: Mitsch, W.J., L. Zhang, A.E. Altor (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2004, pp. 137-148.

ix Fink, D.F. 2006. Analysis of Phragmites australis haplotypes at the ORWRP based on morphological characteristics. In: Mitsch, W.J., L. Zhang, and C. Tuttle, K. Jones (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2005, pp. 127-132.

Fink, D.F. and W.J. Mitsch. 2006. The effect of the removal of hydrologic pulsing on a river-diversion riparian wetland. In: Mitsch, W.J., L. Zhang, and C. Tuttle, K. Jones (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2005, pp. 133-148.

Mitsch, W.J., D.F. Fink, L. Zhang, 2002. Net primary productivity of macrophyte communities after eight growing seasons in experimental planted and unplanted marshes. In: Mitsch, W.J., L. Zhang (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2001, pp. 43-48.

Mitsch, W.J., L. Zhang, N. Dillon, and D.F. Fink. 2004. Biogeochemical patterns of created riparian wetlands: Tenth year results (2003). In: Mitsch, W.J., L. Zhang, and C. Tuttle (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2003, pp. 59-68.

Zhang, L., W.J. Mitsch, D.F. Fink. 2004. Hydrology, water quality, and restoration potential for the Upper Big Darby Creek, Central Ohio. In: Mitsch, W.J., L. Zhang, C. Tuttle (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2003, pp. 207-218.

Zhang, L., W.J. Mitsch, C.L. Tuttle, and D.F. Fink. 2006. Water budgets of the two Olentangy River experimental wetlands in 2005. In: Mitsch, W.J., L. Zhang, and C. Tuttle, K. Jones (eds.), Olentangy River Wetland Research Park at the Ohio State University: Annual Report 2005, pp. 25-36.

Technical Reports

Mitsch, W.J., and D.F. Fink. 2001. Wetlands for Controlling Nonpoint Source Pollution from Agriculture. Indian Lake Wetland Demonstration Project Logan County, OH. Final Report. 18 pp.. Indian Lake Watershed Project, Bellfontaine, OH.

FIELDS OF STUDY

Major Field: Environmental Sciences

Specializations: Wetland ecology and restoration Systems ecology and ecosystem modeling

x

TABLE OF CONTENTS

Abstract………………………………………………………………………………….. ii

Dedication………………………………………………………………………………..vi

Acknowledgements……………………………………………………………………...vii

Vita……………………………………………………………………………………...viii

List of Figures…………………………………………………………………………...xv

List of Tables………………………………………………………………………….... xx

1. INTRODUCTION…………………………………………………………………… 1

1.1 Rationale and significance…………..……………………………………….. ...2 1.2 Creation, restoration, and mitigation……………………………………….. ….2 1.3 Ecosystem modeling…………………………………………………………… 4 1.4 A whole-ecosystem approach………………………………………………….. 4 1.5 Goals and objectives………………...…………………………………...…….. 5 1.6 Literature cited……………………………………………………………….…8

2. NUTRIENT BIOGEOCHEMISTRYAND MACROPHYTE ASEMBLAGES IN A CREATED WETLAND……………………………………………….……… ……12

2.1 Abstract……………………………………………………………………..…12 2.2 Introduction……………………………………………………………..……..13 2.3 Methods…………………………………………………………………….… 16 2.3.1 Site description……………………………………………………….… 16 2.3.2 Hydroperiod.……………………………………………………………. 17 2.3.3 Water quality………………………………………………………..….. 18 2.3.4 Statistical methods……………………………………………………… 19 2.4 Results……………………………………………………………………… ...20 2.4.1 Hydrologic loading and hydroperiod ………….………………………..20 2.4.2 Vegetation…………………………………………………………….…21 2.4.3 Nutrients and ………………………………………………... 22 2.5 Discussion……………………………………………………………….…….27 2.5.1 Water quality dynamics…………………………………………..…….. 27 2.5.2 Spatial patterns…………………………………………………..……... 28

xi 2.5.3 Nutrient loading and retention rates……………………………………. 29 2.5.4 Vegetation dynamics in a diversion wetland…………………………… 30 2.6 Conclusions………………………………………………………………...… 30 2.7 Acknowledgements…………………………………………………..………..31 2.8 Literature cited…………………………………………………….….……….32

3. THE EFFECT OF REMOVING HYDROLOGIC PULSING ON A RIVER DIVERSION RIPARIAN WETLAND…………………………….…….... 53

3.1 Abstract………………………………………………………………………..53 3.2 Introduction………………………………………………………………..…..54 3.2.1 Goals and objectives……………………………………………………. 56 3.3 Methods…………………………………………...………………………….. 56 3.3.1 Site description…………………………………….…………………… 56 3.3.2 Hydrologic conditions.…………………………………...…………….. 57 3.3.3 Water quality……………………………………………………..…….. 58 3.3.4 Chemical analysis………………………………………………………. 59 3.3.5 Avian use……………………….………………………………………. 60 3.3.6 Statistical methods……………………………………………………… 61 3.4 Results……………………………………………………..…………………….62 3.4.1 Hydrologic loading and hydroperiod…………………….………….…..62 3.4.2 Nutrient loading and retention rates……………………………….…… 63 3.4.3 Seasonal and spatial nutrient dynamics………………………………… 64 3.4.4 Avian use………………………….……………………………………. 68 3.5 Discussion……………………………………...………………………………..69 3.5.1 The effect of pulsing……………………….…………………………… 69 3.5.2 Comparison to other river diversions………….……………………….. 70 3.5.3 Spatial patterns of nitrogen removal.……………..………………….….71 3.5.4 Management implications…………………………………….……...… 72 3.5.5 Avian effects………………………………………………………….. ...73 3.5.6 Conclusions………………………………………………………...… ...74 3.6 Acknowledgements……………………………………………………...…… 75 3.7 Literature cited…………………………………………………………….…..76

4. THE EFFECT OF REMOVINGAND REINTRODUCING HYDROLOGIC PULSES ON PRODUCTIVITY OF A RIVER-DIVERSION WETLAND……… 100

4.1 Abstract…………………………………….………………………………...100 4.2 Introduction…………………………………….…………………………….101 4.2.1 Goals and objectives……………………...…………………………… 102 4.3 Methods……………………………………………...…………………...… …103 4.3.1 Site description………………………………...……………………. ...103 4.3.2 Hydroperiod…………………………………………..…………….…. 104 4.3.3 Macrophyte productivity and diversity……………….…………….. ...105 4.3.4 Statistical methods……………………………………...……………. ..106 4.4 Results……………………………………………………………….……….. .106

xii 4.4.1 Hydrologic loading and hydroperiod…………………………………..106 4.4.2 Macrophyte communities…………………………………….……...... 107 4.4.3 Macrophyte productivity………………………...…………….….… ...107 4.5 Discussion……………………………………………………………………109 4.5.1 The effect of pulsing…………………………………………….…….. 109 4.5.2 Causes for variation in macrophyte productivity……………...……….111 4.5.3 Nutrients and wetland vegetation patterns………...………..………… 115 4.5.4 Management implications…………………………...….…………….. 115 4.6 Acknowledgements…………………………………………………………. 116 4.7 Literature cited…………………………………………………………...…..117

5. USING SHORT-TERM HYDROLOGY DATA TO PREDICT LONG- TERM SUCCESSIONAL DEVELOPMENT IN A RIPARIAN WETLAND..… .141

5.1 Abstract………………………………………………………………………141 5.2 Introduction…………………………………………………………..……... 142 5.2.1 Goals and objectives………………………………………...………… 145 5.2.2 Site description………………………………………………………... 146 5.3 Modelling Methods……………………...……………………………………. 146 5.3.1 Calibration……………………………………………………….……. 148 5.3.2 Sensitivity analysis…………………………………………………..... 148 5.3.3 Integration and modeling techniques…………………………………..149 5.4 Model structure…………………………………………………………… ...... 150 5.4.1 Hydrologic submodel……………………………………………..…....150 5.4.2 Vegetation submodels………………………………………………… 151 5.4.2.1 Sediment accretion subroutine…………………………………. …152 5.4.2.2 Tree growth submodel…………………………………………..… 152 5.4.2.3 Impact of trees on hydroperiod…………………………………… 154 5.4.2.4 Emergent macrophyte growth submodel……………………..…… 155 5.4.3 Ecosystem state determination…………………………………...…… 157 5.5 Results and Discussion………………………………………..…………….. 157 5.5.1 Simulations……………………………………….…………………… 157 5.5.1.1 Simulation 1 – baseline, 2004 hydroperiod…………….…………. 158 5.5.1.2 Simulation 2 – dramatic flood pulsing………………………..… ...158 5.5.1.3 Simulation 3 – pulsing removed, 2005 hydrologic data...……..…. .159 5.5.1.4 Simulation 4 – sediment accretion………………………………..,.160 5.5.1.5 Simulation 5 – beaver ponding…………………...……….……… 161 5.5.1.6 Simulation 6 – 10 years of Olentangy River data…………..……...161 5.5.1.6.1 Ecosystem succession…………………………………… ...161 5.5.1.6.2 Total productivity………………………………………… .163 5.5.2 Wetland succession……………………………………………...……. 164 5.5.3 Sensitivity analysis…………………………………..…….….………. 166 5.5.4 Model accuracy…………………………………...……….…..……… 167 5.5.5 Model limitations…………………………….………………...………168 5.6 Conclusions……………………………………..…………………...……….169 5.7 Acknowledgements………………………………………………………… .170

xiii 5.8 Literature cited……………………………...………………………………..171

References………………………………………………………………...………… ...206 Appendix A – STELLA Code…………………………………………...…………. …222

xiv

LIST OF FIGURES

Figure Page

2.1. Site map of the Wilma H. Schiermeier Olentangy River Wetland Research Park on The Ohio State University campus. The 2.8 ha created oxbow wetland is between the experimental wetlands and the bottomland hardwood forest. Grab-sample locations are marked with a white circle and the seven biomass sampling transects are marked with white lines.... ……..37

2.2. Annual for a created diversion oxbow wetland and Olentangy River in central Ohio, USA for (a) 2003 and (b) 2004…….…………..…. ….....39

2.3. Dominant vegetation communities in the created oxbow wetland in a) 2003 and b) 2004. The area in the 2004 map marked with the dotted lines shows the extent of the spread Xanthium strumarium following prolonged drawdown after vegetation and productivity surveys in this study. The location of biomass sampling transects are marked on Fig. 2.1.……..…... …….41

2.4. Mean total phosphorus (TP) reduction during flood pulses in 2003 (15 of 17 total pulses) and 2004 (7 of 7 total pulses). A positive percentage indicates a net reduction in TP whereas a negative percentage indicates a net export in TP. Error bars show standard error. The number of TP samples measured during each pulse is indicated below each mark…….… …...43

- - 2.5. Kriging diagrams of the a) nitrate+nitrate (NO3 + NO2 ), b) total Kjeldahl nitrogen (TKN), c) total phosphorus (TP), and d) total suspended solids (turbidity-NTU) in a created diversion oxbow wetland in central Ohio. Nutrient sampling locations are shown in Fig. 2.1……………………… ……...45

2.6. Nitrate loading and export during the second major flood event of 2004. Net retention is the difference between the two. The flood event occurred February 3-15, 2004.………………………………………………...…… …….47

xv 3.1 Site map of the Wilma H. Schiermeier Olentangy River Wetland Research Park on The Ohio State University campus. The 2.8 ha created oxbow wetland is between the experimental wetlands and the bottomland hardwood forest. Grab-sample locations are marked with a white circle. The area to the north of the “created oxbow” label is predominantly emergent marsh and the area to the south of the label is predominantly open water.………………………………………………………….……… ..…82

3.2 Inflow (top) and outflow (bottom) control structures in the created oxbow wetland. The Red Field TideflexTM check valve, top, opens via water pressure when the river elevation is higher than the wetland and closes when the river elevation is lower than the wetland water level, and water pressure is removed. Water then flows back to the Olentangy River though an outflow control , bottom.…………………………………. ….…84

3.3 Water inflow rate for the created oxbow wetland between April 2004 and March 2006. April 2004 through March 2005 was a pulsing hydroperiod year while April 2005 through March 2006 was steady-flow hydroperiod year when flood-pulses were removed from the wetland……………….. …...…86

- 3.4 Kriging diagrams of the nitrate + nitrate (NO3 -N), total nitrogen (TN), soluble reactive phosphorus (SPR), total phosphorus (TP), and total suspended solids (TSS) in the created oxbow wetland showing annual mean concentrations for 2004 and 2005. Inflow of river water is on the left of each diagram; outflow is on right. Sampling locations are shown in Fig. 3.1.………………………………………………………………..…… …...88

- 3.5 Kriging diagrams of the nitrate + nitrate (NO3 -N), total nitrogen (TN), soluble reactive phosphorus (SPR), total phosphorus (TP), and total suspended solids (TSS) in the created oxbow wetland during the wet season in 2004 and 2005. Inflow of river water is on the left of each diagram; outflow is on right. Sampling locations are shown in Fig. 3.1..... …....90

- 3.6 Kriging diagrams of the nitrate + nitrate (NO3 -N), total nitrogen (TN), soluble reactive phosphorus (SPR), total phosphorus (TP), and total suspended solids (TSS) in the created oxbow wetland during the dry season in 2004 and 2005. Inflow of river water is on the left of each diagram; outflow is on right. Sampling locations are shown on Fig. 3.1…. …...92

3.7 Number of new avian species-observed per sampling effort in the created oxbow wetland with a pulsing hydroperiod (April 2004 – March 2005) and steady-flow hydroperiod (April 2005 – March 2006)…………………...... 94

xvi 4.1 Site map of the Wilma H. Schiermeier Olentangy River Wetland Research Park on The Ohio State University campus. The 2.8 ha created oxbow wetland is between the experimental wetlands and the bottomland hardwood forest. The locations of the seven biomass sampling transects are marked with white lines………………………………..……………… …..122

4.2 Hydrograph for the created oxbow wetland and the Olentangy River from April 2004 to September 2006.……………………….…………………. ……124

4.3 Macrophyte communities in the created oxbow wetland during pulsing (2004 and 2006) and steady flow (2005) conditions during peak biomass. The area in the 2004 map marked with dotted lines shows the spread of Xanthium strumarium following a prolonged drawdown that occurred after the biomass surveys were completed.………...………………...………. ….…126

4.4 Comparisons of: (a) the community diversity index and percent areal coverage of emergent macrophytes (r2 = 0.97), (b) the community diversity index and the Richardson-Baker flashiness index (r2 = 0.98), and (c) the total macrophyte production of the wetland and the Richardson- Baker flashiness index (r2 = 0.75)……………………………...………….. ….128

4.5 Total net aboveground primary productivity in the created oxbow wetland during pulsing (2004 and 2006) and steady flow (2005) during peak biomass. Labels a and b indicate statistical differences in the mean biomass between years (p = 0.36 between 2004 and 2005; p = 0.05 between 2004 and 2006; p = 0.02 between 2005 and 2006). Error bars indicate standard error……………………….…………………...………. …...130

4.6 Total production (kg yr-1) of the 5 dominant macrophyte communities in the created oxbow wetland in 2004 (pulsing hydrology), 2005 (steady flow hydrology), and 2006 (restored pulsing hydrology). The “mixed macrophyte” community is comprised predominately of: Eleocharis sp., Scirpus americanus, Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis..………………….……….. …...132

4.7 Effects of pulsing on wetland ecosystem services. Arrows indicate the direction of increased service for each process. A bi-directional arrow indicates uncertainty as to the effect of pulsing with the relative size of the arrowhead suggesting the general impact………………………..……….……134

4.8 Pattern of Typha net above ground primary productivity per unit area (NAPP, g dry weight m-2 yr-1) compared to the nitrate + nitrite concentration in the from the inflow, through the emergent marsh, open water, and outflow of the created oxbow wetland.…...……. ……136

xvii

5.1 Hydrarch succession in wetlands with reversals of successional stage caused by allogenic disturbance ………………..…………….………….... ….179

5.2 Odum diagram of interactions between hydraulic forcing functions and potential ecosystem states (open water, mudflat, marsh, and forested wetland). The forcing functions are precipitation, solar inputs, river stage or pumped inflow, and the seed in the wetland. The ecosystem state selected as a result of water depth is indicated by the order in which they are located in the Odum diagram.…………………..……………………..…...181

TM 5.3 STELLA diagram of the hydrologic submodel. Inputs are river (Rstage), rainfall (Pt), and temperature (T). Output is the surface elevation of the water in the wetland (MSL) in both daily and in seasonal time steps. The hydrologic input to the model from the river can be switched from natural river flood pulsing (Rstage and Pipeinflow subroutine) to a predetermined artificial pumping rate (Pump)……………………………………..……….….183

5.4 STELLATM diagram of the woody vegetation submodel. Inputs are the surface elevation of the wetland water (MSLW) and the surface elevation of wetland sediments (MSLL). Output is the diameter (TD) and height (TH) of tree production and the hydraulic lift (H) due to the trees………. ..…..185

5.5 STELLATM diagram of the macrophyte vegetation submodel. Inputs are surface elevation of the wetland water (MSLW), the surface elevation of the wetland sediments (MSLL), the diameter of and trees (TD), and the hydraulic lift (HL). Output is the biomass production of the emergent macrophytes (MacB).………………………………………………………. ....187

5.6 Site map indicating the land surface elevation contours (MSL ft) at which succession simulations were conducted.…………………………………... .…189

5.7 Ecosystem state at the end of five different100 year simulations. Simulation results are a) 2004 hydroperiod, b) 2004 hydroperiod with the variance of the pulse peaks increased by 50% , c) 2005 hydroperiod (steady flow), d) 2004 hydroperiod with sediment accretion, and e) 2004 hydroperiod with the outflow weir height raised by 1 ft (0.3048 m)………. ....191

5.8 Time series of the simulated macrophyte and woody biomass in the created wetland according to a 100-year extrapolation of the stage of the Olentangy River from April 2004 – March 2005 with a sediment submodel. Each graph represents a different elevation contour (Fig. 5.6) within the wetland basin, with a) corresponding to the 723.00 ft MSL contour, b) the 723.43 ft MSL contour, c) the 724.10 ft MSL contour, d) the 724.25 ft MSL contour, e) the 725.00 ft MSL contour, f) the 726.00 ft MSL contour, and g) the 727.00 ft MSL contour.………….……193

xviii

5.9 Time series of simulated macrophyte and woody biomass in the created wetland according to a 100-year extrapolation of the last 10 years of Olentangy River Stage (1996-2005). Each graph represents a different elevation contour (Fig. 5.6) within the wetland basin, with a) corresponding to the 723.00 ft MSL contour, b) the 723.43 ft MSL contour, c) the 724.10 ft MSL contour, d) the 724.25 ft MSL contour, e) the 725.00 ft MSL contour, f) the 726.00 ft MSL contour, and g) the 727.00 ft MSL contour.……………………………………………..……... ….195

5.10 Comparison of simulated and actual elevation of surface waters from April 2003 – March 2006. The model was calibrated for April 2004 – March 2005 and validated for April 2003 – March 2004 and for April 2005 – March 2006………………………………………………………………. ……197

xix

LIST of TABLES

Table Page

2.1 Frequency and duration of flood pulses in a created oxbow wetland on the Olentangy River in central Ohio, USA …………………………………….…...48

2.2 Estimated net aboveground primary productivity, based on peak biomass, in the created oxbow in 2004 for the 11 most dominant emergent macrophytes. The “mixed wetland vegetation” in this table refers to grouped FACW species that individually made up less than 0.1% of the total macrophyte biomass in the wetland….………………………...…….. …...49

2.3 Mean nutrient concentrations and turbidity [SRP = soluble reactive - - phosphorus, TP = total phosphorus, NO3 + NO2 = nitrate and nitrite, TN = total nitrogen, and TSS = total suspended solids] in the oxbow wetland created at the Olentangy River Wetland Research Park, 2003-2004, when - flooded river water is flowing through the wetland. TN = TKN + NO3 + - NO2 = (as N)………………………………………………………….. ..………50

- 2.4 Annual mean loading and removal of nitrate-nitrite (NO3 ), total nitrogen (TN), soluble reactive phosphorus (SRP), and total phosphorus (TP) during 2004 in the river diversion oxbow (ave ± std. error). Rates are in g- N m-2 yr-1 or g-P m-2 yr-1 as is appropriate. The mean loading, export, and the retention of the nutrients during the eight flood pulses in the mitigation oxbow wetland in 2004 are also shown. Rates and the mass retention are calculated according to the number of actual days of flow; note that the duration of the inflow and outflow are different…………….…………….. …...51

2.5 Annual and per pulse removal of nitrate (as N) and total phosphorus in selected created river diversion and riparian wetlands.…………………… ……52

3.1 Inflow and outflow of the oxbow wetland in April 2004 – March 2006. Hydraulic retention time (HRT) was calculated only for days in which there was no inflow or outflow. The Richardson-Baker flashiness index (R-B index) for the created oxbow wetland was calculated on a monthly basis for pulsing and steady-flow hydroperiods. Wet season is defined as November through June; dry season is defined as July through October… ……95

xx - 3.2 Annual mean loading and retention of nitrate-nitrite (NO3 -N), total nitrogen (TN), soluble reactive phosphorus (SRP), and total phosphorus (TP) during pulsing hydroperiod (2004) and steady-flow (2005) years in the river diversion oxbow (ave ± std. error). Rates are in g-N m-2 yr-1 or g- P m-2 yr-1 as is appropriate. Rates and the mass retention during the pulsing year are calculated according to the number of actual days of flow; note that the duration of the inflow and outflow are different.……..……. …….96

3.3 Nutrient and suspended sediment concentrations (ave ± std error (# samples) in the created oxbow wetland at the Olentangy River Wetland Research Park during pulsing (April 2003 – March 2005) and non-pulsing (April 2005 – March 2006) conditions when flooded river water is flowing through the wetland (SRP = soluble reactive phosphorus, TP = total - - phosphorus, NO3 + NO2 = nitrate and nitrite, TN = total nitrogen, and TSS = total suspended solids)……...……………………………………. ……..97

3.4 Bird species observed utilizing the created oxbow wetland with a pulsing hydroperiod, April 2004 – March 2005, and with a steady-flow hydroperiod, April 2005 – March 2006………...………………………….. …...98

3.5 Annual and per pulse removal of nitrate (as N) and total phosphorus in selected created river wetlands.………………………………...………… ...…..99

4.1 Inflow, outflow, mean hydraulic retention time (HRT; ave ± std error), and water depth (ave ± std dev) of the mitigation oxbow wetland in April 2004 – Sept 2006. Units of inflow and outflow are cubic meters of water flux per square meter of wetland area (2.8 ha). HRT was only calculated for days in which there was inflow or outflow. The Richardson-Baker (R-B index) flashiness index for the created oxbow wetland was calculated on a monthly basis for the pulsing and steady-flow hydrologic conditions. The Wet Season is defined as November through June; the Dry Season is defined as July through October. *The second pulsing year (2006) only encompasses a partial year (170 days, April 2006 to September 2006)..….. ….137

4.2 Percent areal coverage and the Community Diversity Index (CDI) of the 5 dominant macrophyte communities in the created oxbow wetland in 2004 (pulsing hydrology), 2005 (steady flow hydrology), and 2006 (restored pulsing hydrology). The “Mixed macrophyte” community is comprised predominately of: Scirpus americanus, Eleocharis sp., Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis. NAPP is calculated for the area where macrophytes actually grew and not for the area of the entire oxbow wetland. CDI = Σ (Ciln(Ci)). Where Ci is the present cover of community “i” and N is the number of macrophyte communities (Mitsch et al., 2005a).…………………………. ..…138

xxi 4.3 Percent of the total biomass comprised by each of the 5 dominant macrophyte communities in the created oxbow wetland in 2004 (pulsing hydrology), 2005 (steady flow hydrology), and 2006 (restored pulsing hydrology). The “mixed. macrophyte” community is comprised predominately of: Eleocharis sp., Scirpus americanus, Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis....…………………………………………………………………..….139

4.4 Net aboveground primary productivity per unit area of vegetation (NOT total production) for the 5 dominant macrophyte communities in the created oxbow wetland in 2004 (pulsing hydroperiod), 2005 (steady-flow hydroperiod), and 2006 (restored pulsing hydroperiod). Superscripts denote significance between years for each macrophyte community (p < 0.05). The “mixed macrophyte” community is comprised predominately of: Eleocharis sp., Scirpus americanus, Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis..…………… …140

5.1 Four potential ecosystem successional stages predicted by the simulation model.………………………………………………………………………. …198

5.2 Model parameters, definitions, values, and sources for the created oxbow wetland hydrologic submodel and sediment accretion sub-routine………... …199

5.3 Model parameters, definitions, values, and sources for the created oxbow wetland tree submodel.……………………………………………………. …..200

5.4 Tree species growth coefficients that can be used in the tree growth model. Coefficients were calculated according to Botkin (1993) unless otherwise noted. GT, β0, β1, b2, b3 and DegD for a given species were calculated using age, height, and diameter parameters, which were taken from Petrides (1988), Sargent (1933), and Jenkins et al. (2003). Values of TOD were taken from Pearlstine et al. (1985) or calculated using from Pretides (1988) and Sargent (1933) as marked. The species used in the model presented here is Populus deltoides.……………………………...………... ….201

5.5 Model parameters, definitions, values, and sources for the created oxbow wetland macrophyte submodel…………………………………………….. ….202

5.6 Growth characteristics and parameters for functional emergent macrophyte groups used in the macrophyte growth sub-model (from Ellison and Bedford, 1993; model calibration). The functional group used in the model presented here is a combination of groups 1 and 3 from Boutin and Keddy (1993).………………………………………………..… ....203

xxii 5.7 Predicted basal area and net above-ground primary productivity (NAPP) per square meter of each elevation zone at the end of 8 (present year) and 100 years given a 100 year extrapolation of the past 10 years of Olentangy River flow. “Total production” refers to the entire wetland basin (multiplying the results from the unit model by the area of the wetland within the given contour band)…………………………………………….. ….204

5.8 Comparison of the state variable sensitivities (Sx) to different model parameters…..……………………………………………………………..…...205

xxiii

CHAPTER 1

INTRODUCTION

The most productive ecosystems are generally those that receive pulses of energy from external sources (e.g., tides, river , pulses of runoff, upwelling) in addition to solar energy (Nixon, 1988; W.E. Odum et al., 1995, Odum, 2000). At the same time, they make the system more “open” to exchange of elements with adjacent systems, which not only may enhance productivity but may also contribute to the stability of the ecosystem. This concept of stability of ecosystems in the face of continued pulses was termed pulse-stability by E.P. Odum (1983, 1995). A flood-pulse hydroperiod (the periodic flooding from an adjacent river or lake) that was once common to riverine wetlands in the Midwest United States (Baker et al., 2004), has been shown to increase nutrient uptake and plant productivity in riparian wetlands around the world (Mitsch and

Ewel 1979, Junk et al., 1989; Tockner et al., 2000). There may be the situation in Ohio as well because many of the river waters have high nutrient loads that can have a

“fertilizer effect” on wetland vegetation (Spink et al., 1998; Mitsch and Gosselink, 2000;

Gusewell et al., 2003; Gathumbi et al., 2005). A flood-pulse hydroperiod , however, can be a stress as well as a subsidy. When inundation is prolonged, flooding may impede plant growth and nutrient uptake (van der Valk and Davis, 1978; Mitsch and Rust, 1984;

Kozlowski, 2002).

1

1.1 Rationale and Significance

As river diversions become an increasingly popular design method for the

ecological engineering of watersheds and deltas (e.g., Reyes et al., 2000), it is important

to evaluate these ecosystems in enough detail to determine how well they function during

infrequent but important hydrologic events such as floods, rainstorms, and ice

melts. At these times the nutrient retaining functions, and storage capacity of a wetland

can be greatly diminished (Novitzki 1982; Raisen and Mitchel, 1995; Raisen et al., 1997).

Some studies have showed that certain designs of created wetlands can compromise

retention ability (Wong and Somes, 1995; Fink and Mitsch, 2004). In addition to the

water quality improvement functions of river diversion wetlands, increasing the

connectivity between and can have a marked impact upon floral and

faunal communities (Brown, 1981; Waters and Shay, 1992; Toner and Keddy, 1997;

Newman et al., 1998; Giovanni and Da Motta Marques, 1998; Tanner et al., 1999;

Casanova and Brock, 2000; Henry et al., 2002; Kellogg et al., 2003) and directly affect chemical and physical processes governing nutrient and suspended solid dynamics within wetlands (Knight et al., 1987; Kadlec, 1994; Boustany et al., 1997; Lane et al., 1999,

2003; Blahnik and Day, 2000; Mitsch and Gosselink, 2000; Spieles and Mitsch, 2000).

1.2 Creation, Restoration, and Mitigation

When we lose wetlands, we lose their ability to provide valuable ecosystem services such as water quality improvement, sediment retention, flood prevention, and habitat for a variety of animals and plants (Mitsch and Gosselink, 2000). The wetland

2 mitigation policy is supposed prevent the loss of these ecosytesm services by requiring that lost wetlands be replaced by newly created or restored wetlands (US EPA, 1990;

Reinartz and Warne, 1993; Mitsch and Gosselink, 2000, Cole and Schafer, 2002).

However, many mitigation, restoration, and creation projects have not been very successful at replacing the functions lost due to the destruction of wetlands for development and agriculture(e.g. Maguire, 1985; Kentula et al., 1992; Mitsch and

Wilson, 1996; Zedler and Callaway, 2000; Cole and Schafer, 2002). One of the reasons for this rate of failure is that there is an inadequate understanding regarding how various ecological, biogeochemical, and hydrological functions of a wetland interact (Cole and

Schafer, 2002). For example, even though such indicators as macrophyte cover, species composition, and hydraulic loading in a wetland are frequently used to determine the legal and ecological success of these wetlands (Reinartz and Warne, 1993; Mitsch et al.,

1998; NRC, 2001; Cole and Schafer, 2002), it is unclear precisely how these features interact within the ecosystem as a whole (Mitsch and Wilson, 1996).

Research is needed to help wetland designers, builder s, and managers create mitigated and restored wetlands in a manner that replaces lost ecosystem functions and not just lost area (Cole and Schafer, 2002). This is important both ecologically and legally as the United State Environmental Protection Agency and the United State Army

Corps of Engineers have stated that mitigation must focus on functional replacement and not simply on area or structure (US EPA, 1990). Mitigation, created and restored wetlands will certainly not perform the intended functions if the landscape position is incorrect and the proper physical structure and characteristic hydrology are lacking (Cole and Schafer, 2002).

3 1.3 Ecosystem Modelling

One way to improve our understanding of how the hydroperiod and plant primary production and succession interact is to develop a simulation model to describe these processes. At this time there are not many, if any, such models that can be readily used by wetland designers to help them create wetlands. For a model to be a useful tool for wetland designers, builders, and managers, it is beneficial for it to be a relatively simple model (so that the cost, time, and effort to acquire the necessary input data is prohibitively high) yet still effectively predict both current ecosystem function and the functional development of a wetland.

Wetland designers may benefit from having simulation models that include both function and form in the same model. The model developed in this study will describe varying developmental scenarios in a created wetland, with a focus on a particular riparian oxbow design. This type of model will potentially suggest variables that can be manipulated to affect ecosystem development. In particular, the depth of water, the accretion of sediment, the role of macrophytes (type, productivity, and cover), and the basin morphology were key features in the modeling effort.

1.4 A whole-ecosystem approach

While there may be no optimal scale for ecosystem experimentation, studies that attempt to link wetland function with structure are often done at inappropriate spatial and temporal scales (Mitsch and Day, 2004). It is not always sound to apply conclusions from small-scale, short-term studies to real-world, full-scale conditions (Ahn and Mitsch,

2002; Mitsch et al., 2005a). Whole ecosystem studies are sometimes criticized because

4 the size, cost, and logistics do not allow for much, if any, replication. These weaknesses, however, are compensated for by the results of whole-ecosystem studies having a lower variability and by being more likely to reveal ecosystem properties that are not apparent in smaller scale experiments (Odum, 1990; Carpenter, 1998; Kemp et al., 2001). For example, even though biotic community structure is often taken as a surrogate for ecosystem function (Kentula et al., 1992), the presence of physical or biological structure may not indicate functional replacement (Campbell et al., 2002). Until our understanding of ecological indicators improves, it is necessary to do in depth whole ecosystem studies to determine what the true functionality of created ecosystems is. Such studies are not often done (Rienartz and Warne, 1993) and there is still much need for improvement in the building of wetlands (Mitsch and Wilson, 1996).

1.5 Goals and Objectives

The overall goal for this research was to understand the role of hydrologic pulsing in the functioning of a created riparian river diversion wetland in the Midwestern USA.

Heterogeneity in wetland morphology, hydroperiod, and maturity on wetland biogeochemistry, primary productivity, and successional development were evaluated.

The specific objectives of this dissertation were to:

1. determine the overall efficacy of the wetland at attenuating influent nutrient loads

(Chapter 2);investigate the importance of seasonal hydrologic pulses vs. steady

5 flow conditions on the wetland’s ability to attenuate nitrogen, phosphorus, and

suspended solids and on the use of created riparian wetland by avian species

(Chapter 3);

3. determine the effect of removing and subsequently restoring seasonal hydrologic

pulses on the primary productivity and floral community diversity in the wetland

(Chapter 4); and

4. develop a simulation model that could predict the long-term vegetation patterns in

this created riparian wetland (Chapter 5).

This project involved manipulating pulse and non-pulse fluxes of nutrient-laden waters into adjacent riparian wetlands on a fourth-order river. Nutrient retention was expected to be affected by the properties of the soil water interface, retention time, flood pulse frequency and duration, loading rates, and macrophytes. The macrophytes were also expected to change in response to changes in the wetland hydroperiod. The macrophytes may also prove to have a significant role in nutrient cycling by affecting residence time and by direct nutrient uptake and decay/deposition.

This research was carried out at a created river diversion wetland along the

Olentangy River at the Wilma H. Schiermeier Olentangy River Wetland Research Park at

The Ohio State University, Columbus, OH, USA (Mitsch et al., 1998b; 2005b). The first study (Chapter 2) focused specifically on water quality functions and the development of herbaceous plant communities in this riparian wetland. I hypothesized that the different zones of the wetland would support different biogeochemical functions. I also

6 hypothesized that the greatest reduction of nitrate-nitrogen would occur in the emergent marsh portion of the wetland and the greatest reduction of total suspended solids and total phosphorus would occur in the open water portion of the wetland. All nutrient and sediment dynamics were expected to vary with seasons. This study was intended to provide a baseline for subsequent studies on the effects of variations in hydroperiod on wetland ecosystem function.

A second study (Chapter 3) evaluated the effect of replacing the natural pulsing hydroperiod with a steady inflow of river water provided artificially by submerged pumps. This study evaluated the hypotheses that the wetland would have greater nutrient removal during pulsing conditions and that the wetland would support a greater number of avian species during pulsing conditions.

The third study (Chapter 4) examined the effect of removing and subsequently restoring a pulsing hydroperiod on macrophyte productivity and diversity. It was hypothesized that wetland productivity, areal coverage, and community diversity would decrease when pulsing was removed and increase again when pulsing was restored.

A fourth study (Chapter 5) developed a unit model that was designed to take short-term hydrologic data and make long-term predictions about the development of the created wetland. The primary goal for the model is to accurately predict the successional maturation of this wetland design so that wetland designers and builders will be able to know with relative certainty what functions a similarly designed wetland will support if built in a different location. Simulations investigated the effects of variations in water delivery (both natural and artificial), sediment accretion, and beaver activity.

7 1.6 Literature Cited

Ahn, C. and W.J. Mitsch. 2002. Scaling considerations of mesocosm wetlands in simulating large created freshwater marshes. Ecological Engineering 18:327-342.

Baker, D.B., R..P. Richards, T.T. Loftus, J.W. Kramer. 2004. A new flashiness: Characteristics and applications to Midwestern rivers and . Journal of the American Water Resources Association 40:503-522.

Blahnik, T. and J.W. Day. 2000. The effects of varied hydraulic and nutrient loading rates on water quality and hydrologic distributions in a natural forested treatment wetland. Wetlands 20:48-61.

Boustany, R.G., C.R. Crozier, J.M. Rybczyk, R.R. Twilley. 1997. Denitrification in a south Louisiana wetland forest receiving treated effluent. Wetlands Ecology and Management 4:273-283.

Brown, S.L. 1981. A comparison of the structure, primary productivity, and transpiration of cypress ecosystems in Florida. Ecological Monographs 51:405-415.

Campbell, D.A., C.A. Cole, R.P. Brooks. 2002. A comparison of created and natural wetlands in Pennsylvania, USA. Wetlands Ecology and Management 10:41-49.

Carpenter, S.R. 1998. The need for large-scale experiments to assess and predict the response of ecosystems to perturbation. pp. 287-312. In: Pace, M.L. and P.M. Groffman (eds.), Successes, Limitations, and Frontiers of Ecosystem Science. Springer-Verlag, NY.

Casanova, M.T and M.A. Brock. 2000. How do depth, duration, and frequency of flooding influence the establishment of wetland plant communities. Plant Ecology 147:237-250.

Cole, C.A. and D. Schafer. 2002. Section 404 wetland mitigation and permit success criteria in Pennsylvania, USA, 1986-1999. Environmental Management 30:508-515.

Fink, D.F. and W.J. Mitsch. 2004. Seasonal and storm event nutrient removal by a created wetland in an agricultural watershed. Ecological Engineering 23:313-325.

Gathumbi, S.M., P.J. Bohlen, D.A. Graetz. 2005. Nutrient enrichment of wetland vegetation and sediments in subtropical pastures. Soil Science Society of America Journal 69:539-548.

Giovanni, S.GT. and D.M.L. Da Motta Marques. 1998. Establishment of three emergent macrophytes under different water regimes. Water Science and Technology 40:233- 240.

8 Gusewell, S., U. Bollens, P. Ryser, F. Klotzli. 2003. Contrasting effects of nitrogen, phosphorus and water regime on first- and second-year growth of 16 wetland plant species. Functional Ecology 17:754-765.

Henry, C.P., C. Amoros, N. Roset. 2002. Restoration ecology of riverine wetlands: A 5- year post-operation survey on the Rhône River, France. Ecological Engineering 18: 543-554.

Junk, W. J. 1999. The of large rivers: Learning from the tropics. Archiv für Hydrobiologie 115: 261–280.

Kadlec, R. H. 1994. Detention and mixing in free water wetlands. Ecological Engineering 3:1–36.

Kellogg, C.H., S.D Bridgham, S.A. Leight. 2003. Effects of water level , shade, and time on germination and growth of freshwater marsh plants along a simulated successional gradient. Journal of Ecology 91:274-282.

Kemp, W.M., J.E. Peterson, R.H. Gardner. 2001. Scale-dependence and the problem of extrapolation: Implications for experimental and natural coastal ecosystems. pp. 3-57 In: Gardner, R.H., W.M. Kemp, V.S. Kennedy, and J. Peterson (eds), Scaling Relationships in Experimental Ecology. Columbia University Press, NY.

Kentula, M.E., Brooks, R.B., Gwin, S.E., Holland, C.C., Sherman, A.D. Sifneos, J.C. 1992. An Approach To Improving Decision Making in Wetland Restoration and Creation Island Press, Washington, D.C.

Knight, R. L., T. W. McKim, H. R. Kohl. 1987. Performance of a natural wetland treatment system for wastewater management. Journal of Control Federation 59:746–754.

Kozlowski, T.T. 2002. Physiological-ecological impacts of flooding on riparian forest ecosystems. Wetlands 22:550-561.

Lane, R.R., J.W. Day, B. Thibodeaux. 1999. Water quality analysis of a freshwater diversion at Caernarvon, Louisiana. 22:327–336.

Maguire, C.E. 1985. Wetland replacement evaluation. Contract No. DACW-65-85-D- 0068. U.S. Army Corps of Engineers, Norfolk District, Virginia.

Mitsch, W.J. and K.C. Ewel. 1979. Comparative biomass and growth of cypress in Florida wetlands. American Midland Naturalist 101:417-426.

Mitsch, W.J. and W.G. Rust. 1984. Tree growth responses to flooding in a bottomland forest in northeastern Illinois. Forest Science 30:499-510.

9 Mitsch, W.J. and R.F. Wilson. 1996. Improving the success of wetland creation and restoration with know-how, time, and self-design. Ecological Applications 6:77-83.

Mitsch, W.J., S. Johnson, M. Liptak. 1998. Planting and planting success of the new mitigation wetland at the Olentangy River Wetland Research Park in 1997. pp. 205- 210. In: Mitsch, W.J. and V. Bouchard (eds). Olentangy River Wetland Research Park Annual Report 1997, Ohio State University, Columbus, OH.

Mitsch, W.J and J.G. Gosselink. 2000. Wetlands, 3rd ed. John Wiley and Sons, NY.

Mitsch, W.J. and J.W. Day, Jr. 2004. Thinking big with whole ecosystem studies and ecosystem restoration—A legacy of H.T. Odum. Ecological Modelling 178:133-155.

Mitsch, W.J., J.W. Day, L. Zhang, R.R. Lane. 2005a. Nitrate-nitrogen retention in wetlands in the Mississippi River Basin. Ecological Engineering 24:267-278.

Mitsch, W.J., L. Zhang, C.J. Anderson, A. Altor, M. Hernandez. 2005b. Creating riverine wetlands: Ecological succession, nutrient retention, and pulsing effects. Ecological Engineering 25:510-527.

National Research Council. 1992. Restoration of Aquatic Ecosystems. National Academy Press, Washington, DC. 552 pp.

Newman, S., J. Schuette, J.B. Grace, K. Ritchey, T. Fontaine, K.R. Reddy, M. Pietrucha. 1998. Factors influencing cattail abundance in the northern everglades. Aquatic Botany 60:265-280.

Nixon, S.W., 1988. Physical energy inputs and the comparative ecology of lake and marine ecosystems. and Oceanography 33:1005–1025.

Novitzki, R.P. 1982. Hydrology of Wisconsin Wetlands: Wisconsin Geological and Natural History Survey Information Circular 40, Madison, WI, 22 p.

Odum, H.T. 1983. Systems Ecology – An Introduction. John Wiley and Sons, New York. Pp. 644.

Odum, E.P., 1990. Field experimental tests of ecosystem-level hypotheses. Trends in Ecological Evolution 5:204–205.

Odum W.E., E.P. Odum. H.T. Odum. 1995. Nature’s pulsing paradigm. Estuaries 18:547–555.

Odum, E. P. 2000. Tidal marshes as outwelling/pulsing systems. In M. P. Weinstein and D. A. Kreeger, eds. International Symposium: Concepts and Controversies in Tidal Marsh Ecology. Kluwer Academic Publishers, Dordrecht.

10

Raisen, G.W. and D.S. Mitchel. 1995. The use of wetlands for the control of non-point source pollution. Water Science and Technology 32:177-186.

Raisen, G.W., D.S. Mitchel, R.L. Croome. 1997. The effectiveness of a small constructed wetland in meliorating diffuse nutrient loadings from an Australian rural catchment. Ecological Engineering 9:19-36.

Reinartz, J.A. and E.L. Warne. 1993. Development of vegetation in small created wetlands in southeastern Wisconsin. Wetlands 13:153-164.

Reyes E, M.L. White, J.F. Martin, G.P. Kemp, J.W. Day, A. Aravamuthan. 2000. Landscape modeling of coastal habitat change in the Mississippi Delta. Ecology 81, 2331–2349.

Spieles, D.J. and W.J. Mitsch. 2000. The effects of season and hydrologic and chemical loading on nitrate retention in constructed wetlands: a comparison of low- and high- nutrient riverine systems. Ecological Engineering 14:77-91.

Spink, A., R.E. Sparks, M. van Oorschot, T.W. Verhoenven. 1998. Nutrient dynamics of large river floodplains. Regulated rivers: Research and Management 14:203-216.

Tanner, C.C., J. D’Eugenio, G.B. McBride, J.P.S. Sukias, K. Thompson. 1999. Effect of water level fluctuation on nitrogen removal from constructed wetland mesocosms. Ecological Engineering 12:67-92.

Tockner , K., F. Malard, J.V. Ward. 2000. An extension of the flood pulse concept. Hydrologic Processes 14:2861-2883.

Toner, M. and P. Keddy. 1997. River hydrology and riparian wetlands: A predictive model for ecological assembly. Ecological Applications 7:236-246.

U.S. Environmental Protection Agency. 1983. Methods for Chemical Analysis of Water and Wastes. 600/4-79-020, U.S. Environmental Protection Agency, Cincinnati, OH.

van der Valk, A.G. and C.B. Davis. 1978. The role of seed banks in the vegetation dynamics of prairie glacial marshes. Ecology. 59:322-335.

Waters, I. and J.M Shay. 1992. Effect of water depth on population parameters of Typha glauca stand. Canadian Journal of Botany 70:349-351.

Wong, T.H.F., and N.L.G. Somes. 1995. A stochastic approach to designing wetlands for pollution control. Water Science and Technology 32:145-151.

Zedler, J.B. and J.C. Callaway. 2000. Evaluating the progress of engineered tidal wetlands. Ecological Engineering 15:211-225.

11

CHAPTER 2

NUTRIENT BIOGEOCHEMISTRYAND MACROPHYTE ASEMBLAGES IN A

CREATED WETLAND

2.1 Abstract

A better understanding of the function of riparian wetlands is needed. In this study, hydrological, successional, and water-quality dynamics are documented for a whole-ecosystem study involving a 3-ha created riparian wetland at the Schiermeier

Olentangy River Wetland Research Park at The Ohio State University in Columbus, Ohio

USA during 2003 and 2004. This wetland typically receives seven or eight natural weeklong flood pulses each year from the Olentangy River. Of 21 species planted in

1997, only Scirpus americanus and Juncus effusus remained as important macrophyte species during the study seven years after planting. The dominant macrophyte in most of the wetland was Typha spp. (angustifolia and latifolia). Mean removal per flood pulse for nitrate-nitrite, total Kjeldahl nitrogen (TKN), soluble reactive phosphorus (SRP), and total phosphorus (TP) was 0.71 g-N m-2, 0.92 g-N m-2, 0.016 g-P m-2, and 0.08 g-P m-2

- respectively. The annual reductions of N-NO3 , TN, P-SRP, and TP were 74%, 41%,

- 46%, and 31% by mass. A greater attenuation of NO3 and TP occurred in the emergent marsh section of the wetland than the open water section. TKN concentration increased

12 through the emergent marsh and subsequently decreased through the open water section.

The created oxbow successfully removed nitrate and phosphorus and also had an acceptable assemblage and coverage of macrophytes. It is recommended that similar diversion wetlands be created in other locations to examine their function under different climatic and hydrological conditions.

Keywords: Created wetlands; Ohio River Basin; Olentangy River Wetland Research

Park; pulsing; river diversion; Typha spp.; wetland function; wetland restoration.

2.2 Introduction

The National Research Council (1992) called for the creation and restoration of 4 million ha of wetlands in the United States by 2010. Mitsch et al. (2001, 2005a) and

Mitsch and Day (2006) have suggested that the restoration of 2 million ha of wetlands is necessary in the Mississippi River basin alone to mitigate the hypoxic zone in downstream coastal waters of the Gulf of Mexico. One of the ideal locations for many of these created and restored wetlands is near rivers where river waters can be diverted mechanically or naturally into a wetland. Riparian wetlands are connected to systems that routinely provide river water and propagules. In turn wetlands provide habitat that serves as a nursery for fish and other aquatic life, improvement of water quality, and flood storage capacity.

River diversion wetlands are wetlands fed primarily by flooding streams, which bring sediments and chemicals into the wetlands along with seasonal floodwaters and for the water to then flow back into the (Mitsch and Day, 2006). Because there are

13 both artificial and natural along major sections of streams, it is often possible to

create such wetlands with minimal construction work, by removing portions of levees to

allow floodwater to enter the wetlands. Such wetlands mimic natural oxbows that capture

floodwater and sediments and slowly release the water back to the river after the flood

passes (Mitsch and Day, 2006).

As river diversions become an increasingly popular method for the ecological

engineering of wetlands in downstream portions of watersheds (e.g., Reyes et al., 2000), it is important to evaluate if this wetland type is also appropriate in the upper portion of watersheds in large catchments. Furthermore, if management nutrient pollution throughout the entire Mississippi watershed is to be attempted, we must have data that detail how specific wetland designs function (Reyes et al., 2000). A few studies have

compared natural and created wetlands in upper and lower portions of large watersheds,

but more study is needed regarding the function of specific river diversion designs in

various climates (e.g., Mitsch et al. 2005a).

There has been a great deal of research on how riparian wetlands function (Mitsch

and Gosselink, 2000; Kozlowski, 2002; Kao et al., 2003) and on the relative success of specific wetland restoration and creation projects (Mitsch and Wilson, 1996; Zedler and

Callaway, 2000; Cole and Schafer, 2002). However, insufficient research has been conducted with regards to the specific design features that lead to optimal functioning of created and restored river diversion wetlands. Also, while many studies have determined the overall efficacy of wetlands, few have analyzed these ecosystems in enough detail to determine if the various zones of the wetland are functioning similarly. In addition, most

14 studies do not explicitly take into account rare but important hydrologic events such as

spring floods, rainstorms, and ice melts. At these times the nutrient retaining functions,

and storage capacity of a wetland can be greatly diminished (Novitzki 1982; Raisen and

Mitchel, 1995; Raisen et al., 1997). Certain designs of created wetlands can compromise

retention ability (Wong and Somes, 1995; Fink and Mitsch, 2004).

The importance of river flood events is increasingly recognized in the field of

restoration ecology and attempts have been made to reconnect rivers with their natural

floodplains (Day et al., 1995; Galat et al., 1998; Hensel et al., 1998; Molles et al., 1998;

Toth et al., 1998; Henry et al., 2002; Mitsch and Day, 2006). In addition to the water

quality merits of river diversion wetlands, increasing the connectivity between rivers and

floodplains can have a marked impact upon floral and faunal communities. One potential

benefit of river diversion wetlands with fluctuating water levels is to create/restore a

mixed habitat of wooded and herbaceous wetland (Toner and Keddy, 1997).

A wetland’s hydroperiod directly affects the nutrient and suspended solid

dynamics within wetlands (Mitsch and Gosselink, 2000). The rate at which a wetland’s

water quality changes is generally acknowledged to be dependant on nutrient

concentrations in the inflow, the chemical form of the nutrient, and water flux (Knight et

al., 1987). For example, maximum efficiency of nitrogen removal occurs at loading rates

below 10 g-N m-2 yr-1 (Lane et al., 2003; Spieles and Mitsch, 2000), when diverted water

is spread out over the largest possible wetland area (Lane et al., 1999; Blahnik and Day,

- 2000), and when the NH3:NO3 ratio is less than 1.0 (Boustany et al., 1997).

Furthermore, because water is unevenly distributed in flow-through wetlands due to

15 different degrees of channelization, microtopography, animal activity, and patterns of vegetative growth, different parts of a diversion wetland will function differently

(Fennessy et al., 1994; Kadlec, 1994).

The objective of this project was to observe the function of a created river diversion wetland in the upper Ohio River basin. This study focused specifically on water quality functions and the development of herbaceous plant communities in this riparian wetland. I hypothesize that the different zones of the wetland will have different biogeochemical functions. Specifically that the greatest attenuation of nitrogen species will occur in the emergent marsh portion of the wetland and that attenuation of total suspended solids and total phosphorus in the open water portion of the wetland will vary with seasons.

2.3 Methods

2.3.1 Site Description

The 3-ha created riparian wetland (referred to here as a created oxbow) examined in this study is located on the of the Olentangy River in Central Ohio at the

Schiermeier Olentangy River Wetland Research Park at The Ohio State University

(Columbus, Ohio, USA; Fig. 2.1). Water enters the oxbow through a Red Field

TideflexTM check valve when the river elevation is higher than the wetland, and flows back to the Olentangy River though an outflow control weir by gravity. The wetland has two significant vegetation zones. The northern half (closest to the inflow) is an emergent marsh, and the southern half (closest to the outflow) is an open water basin. Lack of

16 vegetation in the southern half is likely due to high water conditions during spring, which

may prevent germination of emergent aquatic plants outside of the littoral zone.

2.3.2 Hydroperiod

The wetland’s hydroperiod was determined by measuring the water level of the

created oxbow and the river with staff gauges, and by measuring the flow of water into

and out of the wetland using a Swofer 2100 current meter and an ISCO 730 bubbler

module. A simple mathematical model was developed to describe the inflow based on

the elevation of the river relative to the oxbow water surface and on the shape of the inflow pipe and weir. Inflow into the wetland equaled:

Qin = (2.09)*PipeA*√ (Rstage-MSLW)*43560 (2.1)

Where Qin is the inflow, PipeA is the inundated area of the inflow pipe, Rstage is the

surface water elevation of the river, and MSLW is the surface water elevation of the

wetland. Outflow from the wetland equaled:

1.436 Qout = 10.16*(( MSLW -WL) )*43560 (2.2)

Where Qout is the outflow, and WL is the elevation of the top of the outflow weir (See

Chapter 5 for more details). Daily hydrologic budgets enabled calculation of loading and retention rates (by mass) of the various nutrients.

17

2.3.3 Water Quality

Dawn and dusk inflow and outflow grab samples were taken between October

2002 and October 2004 during these flood events. When the Olentangy River is at a high stage (above 220.9 m MSL), overflow occurs into the diversion wetland from the river. More detailed automatic sampling was conducted during some storm events using

ISCO 6874 autosamplers at the inflow and outflow points of the created oxbow.

Samplers were set to hourly frequencies during high flow periods. This more detailed sampling strategy allowed a comparison of the wetland’s functioning during high and low flow periods. Grab samples (n = 514 over the two water years) were analyzed for nitrate- nitrite, total Kjeldahl nitrogen, soluble reactive phosphorus, total phosphorus, and turbidity.

Manual and auto-sampler grab-samples were split into filtered (0.45 µm) and unfiltered subsamples, and analyzed for nitrogen and phosphorus concentrations.

Nitrate+nitrite, soluble reactive phosphorous (SRP), total Kjeldahl nitrogen (TKN), and total phosphorus (TP) concentrations were measured using Lachet (2000) modifications of standard methods (USEPA 1983, APHA 1998) with a Lachet QuickChem flow- injection analyzer (FIA+ 8000). TP and SRP were analyzed with an ascorbic acid and molybdate color reagent method. TP samples were digested in a block digester using 0.5 ml of a digestion solution made up of H2SO4, K2SO4 and mercuric sulfate and heated for

1 hour at 160oC and 1.5 hours at 380oC. TKN was determined using the salicylate and hypochlorite method. Nitrate+nitrite (hearafter referred to as nitrate or nitrate-N) was

18 determined by the sulfanilamide method after reduction in a cadmium column (Lachet,

2000). Total nitrogen (TN) was estimated to be the sum of TKN and nitrate+nitrite-N.

Total suspended solids were determined using an empirical correlation with turbidity developed for our study site (Harter and Mitsch, 2003). Turbidity was measured in the laboratory using a Hach 2100 N turbidimeter.

Quality control in the laboratory was maintained through the use of matrix duplicates, check standards, matrix spikes, and blanks which were all utilized each time the Lachet was run to verify data precision and accuracy. Matrix duplicates are environmental samples that are divided into two samples in the laboratory and analyzed separately and compared for precision; approximately 5% of the samples were checked with a matrix duplicate. Check standards and blanks are samples of known concentration that are tested and compared to the known value to test for accuracy. If a correlation analysis of the results indicated the test samples were within ± 5% accuracy, then the results were considered acceptable. A matrix spike is an environmental sample to which a known concentration of analyte has been added. Matrix spikes were used to ensure accuracy with 2% of the samples. The spike was taken through the entire analytical procedure and the recovery of the analyte was calculated. Results were expressed as percent recovery of the known amount spiked. The matrix spike was considered acceptable when >95% recovery was achieved.

2.3.4 Statistical Methods

Microsoft Excel and SPSS 11.0 were used to calculate the mean, standard deviation, and correlations between nutrient and other environmental values. When

19 comparing multiple variables, an analysis of variance (ANOVA) was done using

Bonferroni’s method for multiple comparisons. Direct comparisons of single variables

between the first and second water years of the study were done using a homoscedastic,

two-tailed Student’s T-Test for equal means. All tests were conducted at a 95%

confidence interval (α = 0.05).

Interpolated maps of nutrient concentration isobars within the wetland basin were

created with point (punctual) kriging methods using a geostatistical software package

(GS+ Geostatistics for the Environmental Sciences v. 7.0). Prior to interpolation, semi-

variance analysis was done to produce Gaussian (hyperbolic) variogram models of the

autocorrelation present in the data. The active lag distance was set to 40 m (50 % of the

maximum transect length). The lag class distance interval (step size) was selected to

provide the best autocorrelation fit for each individual data set and varied between 2 and

6 m. Anisotropic methods were used as the variation in nutrient concentration was

expected to vary differently between the primary axis of water flow (north-south) and the

axis of lateral water flow (east-west). To account for this variation, an offset angle of 90o with an offset tolerance of 22.5o was used for all geostatistical analyses. Interpolation

values for a specific location were weighed by distance and the degree of autocorrelation

present for that distance. The interpolated isobar maps were created using a uniform x-y

grid that generated 188 points at a space interval of 0.6 m and a search neighborhood of

32 nearest points with no limit on geographical distance.

2.4 Results

2.4.1 Hydrologic loading and hydroperiod

20 From its creation in 1996 through 2004, the 3-ha created oxbow received an

average of seven to eight natural flood pulses each year from the Olentangy River (Table

2.1). Inflow from 1998-2004 averaged 20 ± 4 m yr-1 (or m3 m-2 yr-1; cubic meters of

inflow per square meter of wetland area per year). The Olentangy River typically

provides frequent, short (5-6 days of inflow) flood pulses into the created oxbow. These

pulses typically result in 9-12 days of outflow from the wetland.

During the wet season (November-June) and the dry season (July-October) there

were 165 and 7 days of inflow respectively (Fig. 2.2a). In 2003, the wet season was

atypically dry and the dry season was atypically wet, with 53 days of flow in the wet

season and 41 days of flow in the dry season (Fig. 2.2b). In 2003 and 2004 respectively,

the oxbow received 21 m yr–1 and 27 m yr–1 of water through 17 and 8 independent flood pulses per yr, respectively (Table 2.1). The year 2004 was wet compared to most of the previous years of the wetland’s existence (see previous data in Mitsch and Day, 2006).

Despite differences between the frequency and magnitude of the flood pulses (Fig. 2.2 a,b), the mean duration of flood pulse inflows and outflows were similar. In both years, inflow pulses were 5-6 days in duration and outflow pulses lasted 9-10 days (Table 2.1).

2.3.2 Vegetation

In May 1997, the created oxbow was planted with 6900 rootstocks representing

21 species (Acorus calamus, Alisma plantago-aquatica, Asclepias incarnata,

Cephalanthus occidentalis, Equisetum sp., Iris versicolor, Juncus effusus, Lobelia

cardinalis, Polygonum spp., Pontederia cordata, Potomogeton pectinatus, Sagittaria

latifolia, Saururus cernuus, Schoenoplectus tabernaemontani, Scirpus americanus,

21 Scirpus cyperinus, Scirpus fluviatilis, Sparganium eurycarpum, Spartina pectinata, and

Zizania aquatica (Mitsch et al., 1998). Seventy-five percent of these species are still found growing in the wetland. However, only two of the original planted species,

Scirpus americanus and Juncus effusus, make up a significant portion of the 2003-04 wetland primary productivity (Table 2.2).

The macrophyte species contributing most to productivity are Typha sp.,

Eleocharis sp. and Scirpus americanus (Table 2.2), which together account for 68% of the macrophyte net primary productivity in the wetland. The dominant vegetation communities in the created oxbow are Typha sp., a woody fringe of Salix spp. and

Populus deltoides, a mixed community of Eleocharis sp., Juncus effusus, and Scirpus americanus, and expanding patches of Pontederia cordata (Fig. 2.3). While Typha sp. contributes most to the macrophyte productivity, its proportion of the productivity decreases from 83% at the inflow to 0% at the outflow. Of 105 species of plants identified in the created oxbow in 2003 and 2004, 55 were wetland indicator species

(classified as FACW or OBL). Indicator status was determined using the National

Wetland Indicator List for Region I of USA (Northeast; Reed, 1998).

The changing water coverage in the southern basin of the created oxbow affected the macrophyte cover. As water receded in the late summer, Xanthium strumarium rapidly colonized the exposed mudflat, and became the dominant species on the mudflat until the onset of the first heavy frost.

22

2.4.3 Nutrients and sediments

Although the mean concentration of SRP in the water flowing into the created

oxbow in 2003 (60 ± 1 μg L-1) was almost double the mean concentration in 2004 (33 ± 1

μg L-1; Table 2.3), the mean reduction in SRP concentration was only 11.8% higher.

The pattern of SRP concentration change was also different between the 2003 and 2004.

In 2003 the SRP concentration decreased by the midpoint of the wetland to 13± 1 μg L-1 and then increased to 27± 1 μg L-1 at the outflow. In 2004, there was no decrease in SRP concentration between the inflow and the midpoint followed by a decrease in the second half of the wetland to 19 ± 1 μg L-1.

There was a significant difference in the inflow concentration of TP between

years (p < 0.01). In 2003, the mean inflow concentration as only 92 ± 6 μg L-1 compared

to 203 ± 2 μg L-1 in 2004 (Table 2.3). The percent removal of TP was the same in both years despite the difference in the initial inflow concentration. During early spring high flow periods, the wetland retained phosphorus, but it was a source of phosphorus during some late spring floods and during large thunderstorm events in the drier summer months

(Fig. 2.4). TP concentrations did not decrease informally through the oxbow in either year of the study (Fig. 2.5). TP concentrations increased significantly (p = 0.05) between the midpoint and the outflow of the wetland in 2003. In 2004, TP increased to 230 ± 15

μg L-1 along the northeast bank in the emergent marsh near the inflow of the wetland and

had a significant decrease from the midpoint (36 ± 8 μg L-1) to the outflow

(19 ± 1 μg L-1). There were also differences in the lateral distribution of TP between the

23 two years. In 2003 TP concentrations were fairly uniform east to west across the wetland, where-as in 2004 the east bank had a 20% higher TP concentration than the west bank of the wetland through the emergent marsh.

- - -1 The inflow concentration of NO3 -NO2 in 2003 (4.40 ± 0.04 mg L ) was more than twice as high as the concentration in 2004 (1.81 ± 0.01 mg L-1; Table 2.3). The decrease in nitrate concentration was 13.6% higher in 2004, even though there was a lower average inflow concentration that year. The spatial patterns of nitrate concentrations in the created oxbow were similar between the two years, with was no difference between the concentrations in the and along either shore (Fig. 2.5).

It is unclear if the elevated nitrate concentration seen along the west bank of the created oxbow near the outflow is an artifact of the sampling method or if nitrate was produced within the wetland. Because the sampling method was not continuous, it is possible that the grab sampling did not accurately capture each flood pulse as it traveled through the wetland. During most flood-pulses there was an increase in the nitrate concentration at the inflow as nutrient enriched river water flooded into the created oxbow (Fig. 2.6). As this water moved through a wetland, some amount of the nitrate could be taken up by plants or transformed by denitrifying bacteria in to N2 gas and emitted into the atmosphere (Mitsch and Gosselink, 2000). The nitrate that was not retained or removed flowed through the wetland as a nutrient pulse whose magnitude dropped as it moved through the wetland. If by chance grab sampling for nutrients occurred near the end of this pulse, it would be possible for the peak of the pulse to be near the outflow, making it appear as though nitrate was being produced by the wetland when in reality some amount was simply moving through untransformed.

24 Nitrate could also have been produced within the wetland. Soluble organic

nitrogen from decaying plants and algae can be mineralized under both aerobic and

+ anaerobic pathways through ammonification to form ammonium ions (NH4 ).

Ammonium ions can then be taken up directly by anaerobic organisms, converted to

ammonia (NH3) and lost to the atmosphere via volatilization, or it can be immobilized

through ion exchange with negatively charged soil particles. Ammonium ions can also

be oxidized in the thin oxidized layer that exists at the surface of many wetlands soils by

- Nitrosomonas sp. microorganisms into nitrate (NO2 ) and then by Nitrobacter sp.

- microorganisms into nitrate (NO3 ) in a process called nitrification (Mitsch and

Gosselink, 2000). Nitrification can also occur in the oxidized rhizosphere of plants

where adequate oxygen is often available to convert the ammonium nitrogen to nitrate

nitrogen (Reddy and Graetz, 1988)

The inflow concentration of TN in 2004 was 3.04 ± 0.05 mg L-1 (Table 2.3).

While TN decreased by 24% in 2004, its TKN fraction increased by 200% through the emergent marsh portion of the wetland (Fig 2.5), and then dropped across the open water portion to 0.60 ± 0.01 mg L-1 for an overall increase in TKN of 25.6% in 2004 (Table

2.3), the only year in which it was measured. Concentrations of TKN were higher along the east bank, especially near the end of the emergent marsh area (3.30 ± 0.15 mg L-1),

and lower along the west bank of the oxbow in the emergent zone and at the midpoint

(2.40 ± 0.20 mg L-1). In the open water area, there were no differences in TKN between

the edges and the channel (1.50 ± 0.05 mg L-1). For turbidity, the only lateral difference

was in the open water basin, where both edges were less turbid than the channel.

25 Total suspended solids at the wetland inflow were significantly greater (p = 0.5)

in 2004 (19.5 ± 0.9 NTU) than in 2003 (14.9 ± 1.1 NTU; Table 2.3). By the midpoint of the wetland, there was no difference between years (14.6 ± 2.9 NTU). There was,

however, a significant difference (p = 0.05) between years at the outflow. In 2003 the

total suspended solids dropped 38.9% to 9.1 ± 0.9 NTU. In 2004 there was no significant

drop from the midpoint to the outflow of the wetland (Table 2.3). The most obvious

difference in total suspended solids between the two years is the significant increase (p =

0.01) and subsequent decrease (103 ± 15 NTU) in the middle of the open water portion of

the wetland. Loading and retention rates of the various nutrients were variable throughout

2004 (Table 2.4). The total annual loading rate in 2004 for nitrate-nitrogen, total

nitrogen, soluble reactive phosphorus, and total phosphorus were 32.2 g-N m-2 yr-1, 64.5 g-N m-2 yr-1, 0.48 g-P m-2 yr-1, and 6.1 g-P m-2 yr-1 respectively. Retention rates were 15.4

g-N m-2 yr-1, 32.3 g-N m-2 yr-1, 0.05 g-P m-2 yr-1, and 4.48 g-P m-2 yr-1 respectively for the

same nutrients.

During the eight discrete flood pulses, mean loading for nitrate-nitrogen, total

nitrogen, soluble reactive phosphorus, and total phosphorus was 0.97 g-N m-2 per pulse,

2.54 g-N m-2 per pulse, 0.036 g-P m-2 per pulse, and 0.27 g-P m-2 per pulse. Removal

during the eight flood pulses was 0.71 g-N m-2 per pulse, 0.92 g-N m-2 per pulse, 0.016 g-

P m-2 per pulse, and 0.08 g-P m-2 per pulse respectively (Table 2.4). Overall, there was a

- 74%, 41%, 46%, and 31% reduction in the mass of N-NO3 , TN, P-SRP, and TP

respectively throughout the entire year during eight pulses. Sometimes the oxbow would

receive a flood from the river that was not of sufficient magnitude to create outflow, or

outflow would occur solely as a result of precipitation. These two “incomplete pulse”

26 scenarios are not included in the “during pulse” calculations but they are incorporated in the total annual loading calculation. Also, the “during pulse” calculations are based only on the actual days of flow, whereas the annual calculations are not. Retention rates for

TN, SRP, and TP were all higher early in the year, before the start of the growing season.

Nutrient loading was variable not only between flood pulses, but also within flood pulse as shown in flood data from February 2004 (Fig. 2.4). Peaks in nutrient loading do not necessarily match with the moments of greatest nutrient concentration in the influent or the moments of greatest hydraulic flow. There was no significant correlation for either situation (r2 = 0.25 and r2 = 0.36, respectively).

2.5 Discussion

2.5.1 Water Quality Dynamics

The created oxbow in this study was an effective nutrient sink, especially during the initial mid-winter snowmelt season that is common in this part of Ohio. Nitrate- nitrogen removal was dependent on several hydrologic factors including hydraulic loading rate, water depth, and hydraulic retention time. The created oxbow wetland had a lower removal rate for nitrate during infrequent periods of high flow in the dry season, compared to the wet season flows. The created oxbow’s reduced ability to retain phosphorus during “dry season” flood pulses suggests that perhaps the wetland either reaches its assimilative capacity early in the year, or that the lower water levels and increased carp population combine to produce conditions conducive to the export of suspended solids and phosphorus.

27 Despite large differences in nutrient concentrations from year to year, retention of nutrients (by concentration) was similar each year. For example, total phosphorus retention by concentration was 26% each year despite the fact that mean inflow concentrations in 2004 were more than double the concentrations in 2003. The difference in mean annual concentration of total phosphorus in the inflow is likely the result of a greater percentage of the total samples in 2003 coming from the dry season than in 2004.

More samples were taken in 2004 because there was more rain, and thus more summer flooding in 2003 than in 2004. The difference in the timing and amount of flooding does not necessarily explain the difference in influent nitrate-nitrite levels, which can fluctuate dramatically from year to year in the Olentangy River (Mitsch et al., 2005a).

The increase in turbidity in the open water portion of the oxbow in 2004 coincided with an increased number of common carp (Cyprinus carpio) in the wetland.

Carp swam up the outflow of the oxbow during a large flood pulse from the river in late spring, and took up residence in the wetland.

Much of the fluctuation in total nitrogen is explained by changes in the TKN concentration. Ammonium concentrations within the wetland were negligible

(< 0.05 mg L-1); therefore, since the increase and subsequent decrease in TKN matches vegetation patterns observed in the created oxbow (see below) it is likely that the emergent marsh area contributes organic nitrogen to the water column.

2.5.2 Spatial Patterns

Ninety-two percent of the nitrate-nitrite loss occurs in the emergent marsh in the upper third of the oxbow. This indicates that at this point either the concentration of

28 nitrate-nitrite is too low to be further reduced by wetland processes or that the lack of emergent vegetation in open water areas results in an environment that is not conducive to denitrifying bacteria. The environment may be less suitable because the open water area has fewer plant roots or other plant tissues, and subsequently less organic matter on the benthic substrate for denitrifying bacteria to use as a carbon source. This explanation may also account for the differences between the nitrate concentration in the central channel and the edges of the wetland, as the emergent biomass is significantly greater at the edges. The increase and decrease in TKN and nitrate-nitrite appear to be dependent on inverse conditions. The portions of the wetland that show the greatest decrease in

- NO3 are the areas with the greatest increase in TKN.

There was a greater concentration of TP and SRP along the east bank than along the west bank. Phosphorus can be released from dried soils rich with sorbed phosphorus when these soils are inundated (Olilia et al., 1997). It is likely that higher concentrations of phosphorus in the edge zones were a result of the wetland area expanding and contracting as the oxbow received and discharged water during and following pulses.

2.5.3 Nutrient Loading and Retention Rates

The rates of nitrate-nitrogen and total phosphorus retention are within the ranges

-2 -1 of normal nutrient retention rates in wetlands of 10 - 40 g-NO3-N m yr and 0.5 - 5 g-P m-2 yr-1 reported by Mitsch et al. (2000). The nitrate-nitrogen rate is at the low end of the range predicted by that paper while the phosphorus retention is at the high end. Long- term experience with the adjacent experimental wetlands at this same Ohio location (see

Mitsch et al., 2005b) suggest that nitrate-nitrogen retention has increased or remained

29 steady from year to year while total phosphorus retention has decreased over that time.

The total nitrogen retention rate is also similar to those reported by other studies of river- fed created wetlands (Table 2.5). The created oxbow in this study also retained a greater amount of nitrate-nitrogen and a lesser amount of phosphate-phosphorus when compared to other pulsing systems studied in Australia and South Carolina.

2.5.4 Vegetation dynamics in a diversion wetland

Macrophyte vegetation affects and is affected by water quality in riparian wetlands. In the seven years since the oxbow was created, the extent of the Typha- dominated community has not expanded beyond the upper third of the wetland and this is where most of the nitrate-nitrogen is removed. The vegetation restriction to this location is likely due to the significant reduction of available nutrients in the wetland. By the end of the emergent marsh, nitrate-nitrite concentrations have been reduced to low levels, allowing other plants to better compete with Typha (Koch and Reddy, 1992). Another likely factor is that water levels during spring vegetation emergence are typically quite deep and there is minimal drawdown along the edges of the lower two-thirds of the oxbow. This is not favorable for Typha spp., or any other macrophyte, growth or germination (Keddy and Reznicek. 1986, Squires and van der Valk, 1992). Lastly, there is a long unobstructed east-west fetch on the lower two-thirds of the wetland. It is possible that the Typha community stops where it does more as a result of wind and subsequent wave action than as a result of the nutrient concentrations or water depth.

This question will be better answered as trees on the southwest bank continue to mature and form a more significant windbreak.

30 2.6 Conclusions

The wetland was a net sink for nitrogen and phosphorus during snow-melt, most spring floods, and on an annual basis. The wetland did export nitrogen and phosphorus during some of the flood pulses, most notably during river pulses caused by dry season thunderstorms. Overall, this oxbow design has shown itself to be a success in ecological terms. It would be worth replicating this wetland design in other locations to examine how it functions under a variety of climatic and hydrological conditions.

The created oxbow wetland has developed a diverse and reasonable assemblage of emergent macrophytes although it is dominated by Typha sp. in its upper third closest to the inflow. After nutrients are depleted by this emergent marsh section, more plant diversity occurs in the lower two-thirds of the wetland towards the outlflow. Frequent fluctuations in the water level in the southern section during spring flood events, and the drying out of the southern section into a large mudflat during late summer likely accounts for many of the differences between the two parts of the wetland.

2.6 Acknowledgements

Support provided by U.S. Department of Agriculture NRI CSREES Award 2003-

35102-13518 and a Payne grant from the Ohio Agricultural Research and Development

Center of The Ohio State University. Olentangy River Wetland Research Park publication 07-002. We also thank Maria Hernandez for coordinating the laboratory analysis of nutrient concentrations and Colleen Fink for her help in biomass harvesting.

31 2.7 Literature Cited

American Public Health Association. 1998. Standard Methods for the Analysis of Wastewater, 20 ed., APHA, Washington, DC.

Blahnik, T. and J.W. Day. 2000. The effects of varied hydraulic and nutrient loading rates on water quality and hydrologic distributions in a natural forested treatment wetland. Wetlands 20:48-61.

Boustany, R.G., C.R. Crozier, J.M. Rybczyk, R.R. Twilley. 1997. Denitrification in a south Louisiana wetland forest receiving treated sewage effluent. Wetlands Ecology and Management 4:273-283.

Casey, R.E. and S.J. Klaine. 2001. Nutrient attenuation by a riparian wetland during natural and artificial runoff events. Journal of Environmental Quality 30:1720-1731.

Cole, C.A. and D. Schafer. 2002. Section 404 wetland mitigation and permit success criteria in Pennsylvania, USA, 1986-1999. Env Management 30:508-515.

Day J.W., C.J. Madden, R.R. Twilley, R.F. Shaw, B.A. McKee, M.J. Dagg. 1995. The influence of Atchafalaya River on Fourleague Bay, Louisiana (USA). pp. 151–160. In: Dyer K.R. and R.J. Orth (eds), Changes in Fluxes in Estuaries. Olsen and Olsen, New York,

Fennessy, M.S., C. Brueske, W.J. Mitsch. 1994. Sediment deposition patterns in restored freshwater marshes using sediment traps. Ecological Engineering 3:409-428.

Fink, D.F. and W.J. Mitsch. 2004 Seasonal and storm event nutrient removal by a created wetland in an agricultural watershed. Ecological Engineering 23:313-325.

Galat, D.L., L.H. Frederickson, D.D. Humburg, K.J. Bataille, J.R. Bodie, J. Dohrenwend. 1998. Flooding to restore connectivity of regulated, large-river wetlands (Lower Missouri River). BioScience 48:721–733.

Harter, S.K. and W.J. Mitsch. 2003. Patterns of short-term sedimentation in a freshwater created marsh. Journal of Environmental Quality 32:325-334.

Henry, C.P., C. Amoros, N. Roset. 2002. Restoration ecology of riverine wetlands: A 5- year post-operation survey on the Rhône River, France. Ecological Engineering 18:543-554.

Hensel P.F., J.W. Day, D. Pont, J.N. Day. 1998. Short-term sedimentation dynamics in the Rhone River Delta, France: The importance of riverine pulsing. Estuaries 2:52–65.

32 Kadlec, R. H. 1994. Detention and mixing in free water wetlands. Ecological Engineering 3:1–36.

Kao, J.T., J.E. Titus, W. Zhu. 2003. Differential nitrogen and phosphorus retention by five wetland plant species. Wetlands 23:979-987.

Keddy, P. A., and A. A. Reznicek. 1986. Great Lakes vegetation dynamics: the role of fluctuating water levels and buried seeds. Journal of Great Lakes Research 12:25–36.

Knight, R. L., T. W. McKim, H. R. Kohl. 1987. Performance of a natural wetland treatment system for wastewater management. Journal of Water Pollution Control Federation 59:746–754.

Koch, M.S., and K.R. Reddy. 1992. Distribution of soil and plant nutrients along as a trophic gradient in the Florida Everglades. Journal of Soil Science Society of America 56:1492-1499.

Kozlowski, T.T. 2002. Physiological-ecological impacts of flooding on riparian forest ecosystems. Wetlands 22:550-561.

Lachet Instruments. 2000. Methods Manual. Lachet Instruments, Milwaukee, WI, USA.

Lane, R.R., J.W. Day, and B. Thibodeaux. 1999. Water quality analysis of a freshwater diversion at Caernarvon, Louisiana. Estuaries 22:327–336.

Lane, R.R., H.S. Mashriqui, G.P. Kemp, J.W. Day, J.N. Day, A. Hamilton. 2003. Potential nitrate removal from a river diversion into a Mississippi delta forested wetland. Ecological Engineering 20:237-249.

Mitsch, W.J., J.K. Cronk, X. Wu, R.W. Nairn D.L. Hey. 1995. Phosphorus retention in constructed freshwater riparian marshes. Ecological Applications 5:830-845.

Mitsch, W.J. and R.F. Wilson. 1996. Improving the success of wetland creation and restoration with know-how, time, and self-design. Ecological Applications 6:77-83.

Mitsch, W.J., S. Johnson, M. Liptak. 1998. Planting and planting success of the new mitigation wetland at the Olentangy River Wetland Research Park in 1997. pp. 205- 210. In: Mitsch, W.J. and V. Bouchard (eds). Olentangy River Wetland Research Park Annual Report 1997, Ohio State University, Columbus, OH.

Mitsch, W.J and J.G. Gosselink. 2000. Wetlands, 3rd ed. John Wiley and Sons, New York.

33 Mitsch, W.J., A.J. Horne, R.W. Nairn. 2000. Nitrogen and phosphorus retention in wetlands —Ecological approaches to solving excess nutrient problems. Ecological Engineering 14:1-7.

Mitsch, W.J., J.W. Day, W. Gilliam, P.M. Groffman, D.L. Hey, G.W. Randall, N. Wang. 2001. Reducing nitrogen loading to the Gulf of Mexico from the Mississippi River Basin: strategies to counter a persistent ecological problem. BioScience 51:373-388.

Mitsch, W.J., J.W. Day, L. Zhang, R.R. Lane. 2005a. Nitrate-nitrogen retention in wetlands in the Mississippi River Basin. Ecological Engineering 24:267-278.

Mitsch, W.J., L. Zhang, C.J. Anderson, A. Altor, M. Hernandez. 2005b. Creating riverine wetlands: Ecological succession, nutrient retention, and pulsing effects. Ecological Engineering 25:510-527.

Mitsch, W.J. and J.W. Day, Jr. 2006. Restoration of wetlands in the Mississippi-Ohio- Missouri (MOM) River Basin: Experience and needed research. Ecological Engineering 26:55-69.

Molles M.C., C.S. Crawford, L.M. Ellis, H.M. Valett, C.N. Dahm. 1998. Managed flooding for riparian ecosystem restoration. BioScience 48:748–756.

Nairn, R.W. and W.J. Mitsch. 2000. Phosphorus removal in created wetland ponds receiving river overflow. Ecological Engineering 14:107-126.

National Research Council. 1992. Restoration of Aquatic Ecosystems. National Academy Press, Washington, DC. 552 pp.

Novitzki, R.P. 1982. Hydrology of Wisconsin Wetlands: Wisconsin Geological and Natural History Survey Information Circular 40, Madison, WI, 22 p.

Olila, O.G, K.R. Reddy, D.L. Stites. 1997. Influence of draining on soil phosphorus forms and distribution in a constructed wetland. Ecological Engineering 14:107-126.

Phipps RG and W.G. Crumpton. 1994. Factors affecting nitrogen loss in experimental wetlands with different hydrologic loads. Ecological Engineering 3:399–408.

Raisen, G.W. and D.S. Mitchel. 1995. The use of wetlands for the control of non-point source pollution. Water Science and Technology 32:177-186.

Raisen, G.W., D.S. Mitchel, R.L. Croome. 1997. The effectiveness of a small constructed wetland in meliorating diffuse nutrient loadings from an Australian rural catchment. Ecological Engineering 9:19-36.

34 Reddy, K.R. and D.A. Graetz. 1988. Carbon and nitrogen dynamics in wetland soils. Pp. 307-318. In: D.D. Hook, W.H. McKee, Jr., H.K. Smith, J. Gregory, V.G. Burrel, M.R. DeVoe, R.E. Sojka, S. Gilbert, R.Banks, L.G. Stolzy, C. Brooks, T.D. Mathews, and T.H Shear (eds.) The Ecology and Management of Wetland, Vol. 1: The Ecology of Wetlands. Timber Press, Portland, OR.

Reed, P.B. Jr. 1998. National list of plant species that occur in wetlands: Northeast (Region I). U.S. Fish and Wildlife Service, Washington, DC, Biological Report 88 (26.1).

Reyes E, M.L. White, J.F. Martin, G.P. Kemp, J.W. Day, A. Aravamuthan. 2000. Landscape modeling of coastal habitat change in the Mississippi Delta. Ecology 81:2331–2349.

Spieles, D.J. and W.J. Mitsch. 2000. The effects of season and hydrologic and chemical loading on nitrate retention in constructed wetlands: a comparison of low- and high- nutrient riverine systems. Ecological Engineering 14:77-91.

Squires, L. and A.G. van der Valk. 1992. Water-depth tolerances of the dominant emergent macrophytes of the Delta Marsh, Manitoba. Canadian Journal of Botany 70:1860-1867.

Toner, M. and P. Keddy. 1997. River hydrology and riparian wetlands: A predictive model for ecological assembly. Ecological Applications 7:236-246.

Toth L.A., S.L. Melvin, D.A. Arrington, J. Chamberlain. 1998. Hydrologic manipulations of the channelized Kissimmee River. BioScience 48:757–765.

U.S. Environmental Protection Agency. 1983. Methods for Chemical Analysis of Water and Wastes. 600/4-79-020, U.S. Environmental Protection Agency, Cincinnati, OH.

Wong, T.H.F., and N.L.G. Somes. 1995. A stochastic approach to designing wetlands for stormwater pollution control. Water Science and Technology 32:145-151.

Zedler, J.B. and J.C. Callaway. 2000. Evaluating the progress of engineered tidal wetlands. Ecological Engineering 15:211-225.

35

Figure 2.1 Site map of the Wilma H. Schiermeier Olentangy River Wetland Research

Park on The Ohio State University campus. The 2.8 ha created oxbow wetland is between the experimental wetlands and the bottomland hardwood forest. Grab-sample locations are marked with a white circle and the seven biomass sampling transects are marked with white lines.

36 37

Figure 2.2 Annual hydrograph for a created diversion oxbow wetland and Olentangy

River in central Ohio, USA for (a) 2003 and (b) 2004.

38

39

Figure 2.3 Dominant vegetation communities in the created oxbow wetland in a) 2003

and b) 2004. The area in the 2004 map marked with the dotted lines shows the extent of

the spread Xanthium strumarium following prolonged drawdown after vegetation and productivity surveys in this study. The location of biomass sampling transects are marked on Fig. 2.1.

40

a) 2003 b) 2004

-Open water -Typha spp -Woody fringe -Salix spp. -Populus deltoides -Mixed communities -Eleocharis sp -Scirpus americanus -Juncus effusus -Pontedaria cordata -Xantium strumarium

N

0 200 feet 0 60 meters

Outflow Outflow

41

Figure 2.4 Mean total phosphorus (TP) reduction during flood pulses in 2003 (15 of 17 total pulses) and 2004 (7 of 7 total pulses). A positive percentage indicates a net reduction in TP whereas a negative percentage indicates a net export in TP. Error bars show standard error. The number of TP samples measured during each pulse is indicated below each mark.

42 100% 2003 pulses 75% 2004 pulses

50%

25%

0% 43 -25%

Percent reduction . -50%

-75%

-100% Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Month

- - Figure 2.5 Kriging diagrams of the a) nitrate+nitrate (NO3 + NO2 ), b) total Kjeldahl nitrogen (TKN), c) total phosphorus (TP), and d) total suspended solids (turbidity-NTU) in a created diversion oxbow wetland in central Ohio. Nutrient sampling locations are shown in Fig. 2.1.

44

- -1 -1 a) NO3 mg-N L b) TKN mg-N L

Inflow Inflow Inflow

0 200 feet 2.00 0 60 3.50 2003 2004 2004 meters

N

0.05 0.60

Outflow Outflow Outflow c) TP mg-P L-1 d) Turbidity - NTU

Inflow Inflow Inflow Inflow

0.23 155 2003 2004 2003 2004

0.12 15

Outflow Outflow Outflow Outflow

45

Figure 2.6 Nitrate loading and export during the second major flood event of 2004. Net retention is the difference between the two. The flood event occurred February 3-15,

2004.

46

250

200 Outflow

-1 Inflow yr

-2 150 -N m g 100 m

50

0 2 Feb 2004 6 Feb 2004 10 Feb 2004 16 Feb 2004 Date

47

Mean duration of pulses, days

Pulses per year Inflow Outflow 1996-2002 7-8 5-6 12

2003 17 5-6 10

2004 8 5-6 9

Table 2.1 Frequency and duration of flood pulses in a created oxbow wetland on the

Olentangy River in central Ohio, USA.

48

NPP % of

Species g m-2 yr-1 total

Typha sp. 276 28.0

Leersia oryzoides 66 6.7

Juncus effusus 140 14.2

Verbesina alterniflora 35 3.5

Pontederia cordata 2 0.2

Scirpus americanus 191 19.4

Eleocharis sp. 209 21.2

Phragmites australis 26 2.7

Mixed wetland vegetation 23 2.3

Sparganium eurycarpum 16 1.6

Juncus canadensis 2 0.2

______

Total 986 100

Table 2.2 Estimated net aboveground primary productivity, based on peak biomass, in the

created oxbow in 2004 for the 11 most dominant emergent macrophytes. The “mixed

wetland vegetation” in this table refers to grouped FACW species that individually made up less than 0.1% of the total macrophyte biomass in the wetland.

49

Parameter Inflow Mid-point Outflow % Removal

2003 2004 2003 2004 2003 2004 2003 2004

SRP (μg-P L-1) 60 ± 1 (79)b 33 ± 1 (62)b 13 ± 1 (16)bc 36 ± 8 (6)b 27 ± 1 (97)c 19.0 ± 0.3 (87)c 55.3 43.5

TP (μg-P L-1) 92 ± 6 (8)b 203 ± 2 (77)b 77 ± 5 (7)bc 144 ± 11 (7)bc 68 ± 5 (6)b 150 ± 1 (102)b 25.7 26.3

- -1 b b bc bc b b b b NO3 (mg-N L ) 4.40 ± 0.04 (58) 1.81 ± 0.01 (83) 2.32 ± 0.14 (14) 0.66 ± 0.07 (7) 2.65 ± 0.02 (79) 0.77 ± 0.01 (120) 39.9 57.3

TN (mg-N L-1) - 3.04 ± 0.05 (77) - 3.17 ± 0.35 (7)c - 2.31 ± 0.02 (102)c - 24.0

TSS (mg L-1) 14.9 ± 1.1 (79)b 19.5 ± 0.9 (87)b 14.2 ± 1.9 (16)b 14.6 ± 2.9 (7)bc 9.1 ± 0.4 (99)bc 16.4 ± 0.4 (128)b 38.9b 15.9b 50 Mean ± Standard Error (number of samples) bSignificant difference between years (p < 0.05) cSignificant difference from upstream location (p < 0.05)

- Table 2.3 Mean nutrient concentrations and turbidity [SRP = soluble reactive phosphorus, TP = total phosphorus, NO3

- + NO2 = nitrate and nitrite, TN = total nitrogen, and TSS = total suspended solids] in the oxbow wetland created at the

Olentangy River Wetland Research Park, 2003-2004, when flooded river water is flowing through the wetland. TN =

- - TKN + NO3 + NO2 = (as N).

50 - NO3 -N TN SRP TP

Yearly Mean

Loading Rateg-X m-2 yr-1 32.2 ± 0.2 64.5 ± 0.4 0.48 ± 0.00 6.10 ± 0.04

Export Rate g-X m-2 yr-1 16.8 ± 0.2 32.2 ± 0.3 0.43 ± 0.01 1.62 ± 0.01

Retention

Rate g-X m-2 yr-1 15.4 ± 0.2 32.3 ± 0.2 0.05 ± 0.01 4.48 ± 0.03

Percent (by mass) 48 ± 3 50 ± 4 10 ± 1.0 73 ± 8.0

During Eight Pulses

Loading g-X m-2 pulse-1 0.97 ± 0.11 2.54 ± 0.47 0.036 ± 0.006 0.27 ± 0.07

Export g-X m-2 pulse-1 0.25 ± 0.01 1.34 ± 0.03 0.019 ± 0.004 0.19 ± 0.06

Retention

Total g-X m-2 pulse-1 0.71 ± 0.10 0.92 ± 0.44 0.016 ± 0.003 0.08 ± 0.01

Percent (by mass) 73.9 ± 11.1 40.6 ±6.8 46.8 ± 24.9 31.1 ± 6.1

Unit ‘X’ is N or P, as appropriate

- Table 2.4 Annual mean loading and removal of nitrate-nitrite (NO3 ), total nitrogen (TN), soluble reactive phosphorus (SRP), and total phosphorus (TP) during 2004 in the river diversion oxbow (ave ± std. error). Rates are in g-N m-2 yr-1 or g-P m-2 yr-1 as is

appropriate. The mean loading, export, and the retention of the nutrients during the eight

flood pulses in the mitigation oxbow wetland in 2004 are also shown. Rates and the mass

retention are calculated according to the number of actual days of flow; note that the

duration of the inflow and outflow are different.

51

Annual removal Removal per pulse - -2 -2 - -2 -2 Study Site g NO3 -N m g P m g NO3 -N m g-P m Reference Sustainable retention rates 10 – 40 0.5 – 5 Mitsch et al., 2000 for non-treatment wetlands Riparian oxbow, Columbus, OH 15 4.5 0.71 0.08 This study Riparian wetland, Cooke, 23 2.8 0.353 0.0523 Raisen and Mitchell, 1995 Australia Raisen et al.,1997 Riparian wetland, Cheraw State 0.038 0.067a Casey and Klaine, 2001 Park Golf Course, SC River-fed created wetland, Lake 11-38 1.4-2.9 Phipps and Crumpton, 1994 County, IL Mitsch et al., 1995 52 River-fed created wetland, 58-66 5.2-5.6 Spieles and Mitsch, 2000 Columbus, OH Nairn and Mitsch, 2000 Mississippi River diversion, 31 Lane et al., 1999 Caernarvon, LA Mitsch et al., 2005 a The Cheraw State Park site data are given in phosphate-phosphorus loading.

Table 2.5 Annual and per pulse removal of nitrate (as N) and total phosphorus in selected created river diversion and riparian

wetlands.

52

CHAPTER 3

THE EFFECT OF REMOVING HYDROLOGIC PULSING ON A RIVER

DIVERSION RIPARIAN WETLAND

3.1 Abstract

While hydrologic pulses are key to wetland function and often enhance the productivity and contribute to the stability of ecosystems, the overall effects of these pulses on biogeochemical processes in riparian river diversion wetland ecosystems have not been clearly demonstrated in ecosystem studies. One year of pulsing and one year of steady-flow inputs of nutrient laden river water were compared in a 3-ha created oxbow.

The wetland received 8 flood pulses in 2004 for a total of 27 m yr-1 of inflow and 20 m yr-1 of inflow in the steady flow year 2005. The retention rate was higher for nitrate- nitrogen, total nitrogen, and total phosphorus during the pulsing year then during the steady flow year. The only nutrient species that did not have a difference in retention rate in the two years was soluble reactive phosphorus. There were differences in spatial dynamics of most of the nutrient species in pulsing and non-pulsing years and between wet and dry seasons. In all cases, however, total nitrogen increased through the emergent marsh and then decreased across the open water basin. There was greater avian use of

53 the created oxbow wetland during the pulsing year than in steady flow conditions. The guild of bird species with the greatest difference in presence between the pulsing and steady-flow conditions was shorebirds.

Keywords: Avian use, created wetland, nitrate, phosphorus, wetland design

3.2 Introduction

Flood pulses have both positive and negative effects on the overall biotic function of wetland ecosystems (Day et al.,1977, 1995; Mitsch and Ewel, 1979; Mitsch et al.,

1979; Mitsch and Rust, 1984; Megonigal et al., 1997; McDougal et al., 1997; Galat et al.,

1998; Hein et al., 1999; Day et al., 2000; Reyes et al., 2004; Lane et al., 2004; and Mitsch et al., 2005c). But the overall effects of pulses on biogeochemical and biological processes in these riparian river diversion wetland ecosystems are not clear, nor have they been clearly demonstrated in ecosystem studies (Mitsch and Day, 2006). The role of pulsing in river ecology has evolved from the (Vannote et al.,

1980) to the Flood Pulse Concept described by Junk (1999) and Tockner et al. (2000) which emphasizes the exchange between a river and its floodplain as the primary factor affecting the function of both systems.

Stabilizing water levels or managing them outside the range of historic fluctuations eliminates the dynamic patterns that allow a diversity of wetland species and communities to thrive (Bedford, 1996). Minimizing the fluctuations in Great Lakes fringing wetlands through active management (Wilcox, 1993; Wilcox and Whillans,

1999) has run counter to the maintenance of hemi-marsh conditions that benefit most

54 wetland bird species (Gibbs et al., 1991; Steen et al., 2006). However little literature exists studying whether this same relationship exists in riparian wetlands that have a flood driven pulsing hydroperiod. Understanding the relationships between birds and wetland forcing functions is useful as aquatic birds have been identified as indicators of wetland processes (Fernandez et al., 2005), especially water eutrophication processes

(Rutschke, 1987).

Excessive nutrients have led to significant eutrophication and subsequent hypoxic conditions in coastal waters throughout the developed world. One of the most notable of these conditions is the 20,000 km2 hypoxic zone in the Gulf of Mexico caused by excessive nitrates in the Mississippi River (Rabalais et al., 1996; 1998; 1999; 2002;

Turner et al., 2005). The retention of nutrients, especially nitrogen, in Midwestern USA wetlands could be of key importance as an economical solution to the Gulf of Mexico problem. The creation or restoration of 20,000 km2 of wetlands and riparian ecosystems in the Mississippi River Basin has been recommended as a means to remove nitrogen from the Basin to alleviate the Gulf hypoxic zone (Mitsch et al., 1999; 2001; 2005b). It is necessary to assess the role of these potential wetlands as nutrient sinks and the importance of hydrologic pulses in this function. Nutrient fluxes come as seasonal pulses related to river and runoff discharge, not as fluxes at constant concentrations. If floods flowing into wetlands are synchronized with high concentrations of nitrate, for example, higher nitrogen retention may result (Nixon et al., 1996; Lane et al., 1999). But if hydrologic pulses occur during periods of low nitrate, lower N retention in the wetland may result (Spieles and Mitsch, 2000).

55

3.2.1 Goals and Objectives

This project examines the effects of removing pulse fluxes of nutrient-laden river waters into created riparian diversion wetlands on a fourth-order river in central Ohio,

USA, and replacing these natural flood pulses with artificial steady-flow inputs. Wetland function under both pulsing and steady flow conditions was determined through measurements of avian use and fluvial inflow and outflow of nutrients.

The goal of this study is to investigate the effects of removing seasonal flood pulses on wetlands receiving diverted river water in the Upper Mississippi River Basin.

Pursuant to this goal I had the following objectives:

1. Investigate the effect of removing hydrologic pulsing on the ability of a riparian

wetland to remove nutrients;

2. Determine the effects of removing seasonal flood pulses on avian use in the

wetland.

3.3 Methods

3.3.1 Site Description

The wetland in this study is a created river diversion wetland located at the Wilma

H. Schiermeier Olentangy River Wetland Research Park at The Ohio State University in

Columbus, Ohio, USA (latitude 40.021°N, longitude 83.017°E; Fig. 3.1). A river diversion wetland is a wetland on the adjacent floodplain or behind artificial levees that

56 receive water by pumping or flood flows from the main channel of a river (Mitsch and

Day, 2006). From its creation in 1996 until 2004, the 3-ha created riparian wetland

(referred to here as a created oxbow) received, on average, 7 to 8 natural flood pulses per year from the Olentangy River. Inflow from 1998-2004 averaged 20 ± 4 m yr-1. The

Olentangy River provides frequent short (5-6 days of inflow) flood pulses into the created oxbow which typically result in 9-12 days of outflow from the wetland. Water flows into the northern tip of the wetland through a Red Field TideflexTM check valve when the river elevation is higher than the wetland; the valve closes when the river elevation is lower than the wetland water level and water then flows back to the Olentangy River though an outflow control weir (Fig. 3.2; Fink and Mitsch, in press).

3.3.2 Hydrologic conditions

The 3-ha created oxbow was monitored during one year of flood-pulse conditions,

2004, and then subjected to one year with steady-flow conditions, 2005 (Fig. 3.3).

Pulsing conditions during the first year (April 2004 to March 2005) were created entirely by natural flooding of the river itself. Steady-flow conditions during the second year

(April 2005 – March 2006) were created and controlled by a large submersed bypass- pump on the river intake that eliminated floods and created artificial steady-flow conditions. I attempted to provide a similar volume of water to the wetlands in the non- pulsing year with the pumps as was provided naturally by the river during the pulsing hydroperiod year.

Flows into and out of the wetland were measured using a Swofer 2100 current meter for water velocity and an ISCO 730 bubbler module. A simple mathematical

57 model was developed to describe the inflow based on the relative elevation of the river,

the oxbow water surface elevation, and the flooded cross-sectional area of the inflow

pipe. Outflow was estimated from the stage of water within the wetland and the shape of

the outflow weir (USBR, 1997). A daily hydrologic budget was then developed from

daily readings of river elevation and wetland stage.

To quantify the differences between the pumped steady flow and the natural river

pulsing, the Richardson-Baker flashiness index for streams was applied to the wetland

(Baker et al., 2004).

(1)

Where q is the daily inflow rate. The index measures oscillations in flow relative to total

flow, and as such, provides a useful characterization of hydrologic inputs. This index was calculated on a monthly time step to allow a statistical comparison of flow stability between the pulsing and non-pulsing years.

3.3.3 Water Quality

Dawn and dusk inflow and outflow 500 mL grab samples were taken between

April 2004 and March 2006 on days when there was flow into or out of the created oxbow wetland. During the year with the pulsing hydroperiod (2004), when the

Olentangy River stage reached or exceeded 220.9 m MSL and was higher than the surface water elevation of the wetland, river water was diverted into the oxbow wetland.

More detailed sampling was conducted during some storm events using ISCO 6874 autosamplers at the inflow and outflow points of the created oxbow. Autosamplers

58 collected hourly 500 mL samples during high flow periods. Grab samples were also

taken at 11 locations around the wetland to determine the spatial distribution of nutrients

and suspended sediments (Fig. 3.1). During the pulsing year, these samples were taken

every 4 days during eight flow events; during the steady-flow year, these samples were

taken twice per month. The grab samples (n = 700 over the two water years) were

- analyzed for nitrate + nitrite (NO3 ), total Kjeldahl nitrogen (TKN), soluble reactive phosphorus (SRP), total phosphorus (TP), and total suspended solids (TSS).

3.3.4 Chemical Analysis

Nitrate + nitrite, soluble reactive phosphorous (SRP), total Kjeldahl nitrogen

(TKN), and total phosphorus (TP) were analyzed using Lachet (2000) modifications of

standard methods (USEPA 1983, APHA 1998) with a Lachet QuickChem flow-injection

analyzer (FIA+ 8000). Manual and auto-sampler samples were split into filtered (0.45

µm) and unfiltered subsamples within 48 hours of sampling, and analyzed for appropriate

chemistries. TP and SRP were analyzed with an ascorbic acid and molybdate color

reagent method. TP samples were digested in a block digester using 0.5 mL of a digestion

o solution made up of H2SO4, K2SO4 and mercuric sulfate and heated for 1 hour at 160 C

and 1.5 hours at 380oC. TKN was determined using the salicylate and hypochlorite

method. Nitrate + nitrite was determined by the sulfanilamide method after reduction in

a cadmium column. Throughout this paper references to nitrate or nitrate-N should be

interpreted as references to nitrate + nitrite. Total nitrogen (TN) was estimated to be the

sum of TKN and nitrate + nitrite-N. TKN is a measure of ammonia-N and organic-N.

Measurements of ammonium ion concentration in this wetland are negligible (Fink and

59 Mitsch, in press). Therefore the TKN fraction of the TN can be considered a measure of the organic-N concentration in the wetland surface water. Total suspended solids were determined using an empirical correlation with turbidity developed for the Olentangy

River (Harter and Mitsch, 2003). Turbidity was measured in the laboratory using a Hach

2100 N turbidimeter.

Quality control in the laboratory was maintained through the use of matrix duplicates, check standards, matrix spikes, and blanks which were all utilized each time the Lachet was run to verify data precision and accuracy. Matrix duplicates are environmental samples that are divided into two samples in the laboratory and analyzed separately and compared for precision; approximately 5% of the samples were checked with a matrix duplicate. Check standards and blanks are samples of known concentration that are tested and compared to the known value to test for accuracy. If a correlation analysis of the results indicated the test samples were within ± 5% accuracy, then the results were considered acceptable. A matrix spike is an environmental sample to which a known concentration of analyte has been added. Matrix spikes were used to ensure accuracy with 2% of the samples. The spike was taken through the entire analytical procedure and the recovery of the analyte was calculated. Results were expressed as percent recovery of the known amount spiked. The matrix spike was considered acceptable when >95% recovery was achieved.

3.3.5 Avian use

Avian use of the created oxbow was observed twice-per-month during the pulsing hydroperiod (2004) and steady-flow (2005) years Birds were observed from the same

60 locations as the grab samples were taken (Fig. 3.1), and were only recorded when they

were observed visually. A bird was considered to be using the wetland and recorded only

if it was observed feeding, hunting, sitting, swimming, or walking through the wetland.

Avian species that were only observed flying over the wetland area were not counted.

Because the birds were not tagged, it was not possible to differentiate between

individuals, therefore only the species of bird observed was recorded and not the number

of birds observed.

3.3.6 Statistical Methods

The mean, standard deviation, and correlations between nutrient and other environmental values were calculated using Microsoft Excel and SPSS 11.0. Direct comparisons of single variables between the first and second water years of the study were done using a homoscedastic, two-tailed Student’s T-Test for equal means. All tests were conducted at a 95% confidence interval (α = 0.05).

Interpolated maps of nutrient concentration isobars within the wetland basin were

created with point (punctual) kriging methods using a geostatistical software package

(GS+ Geostatistics for the Environmental Sciences v. 7.0). Prior to interpolation, semi-

variance analysis was done to produce Gaussian (hyperbolic) variogram models of the

autocorrelation present in the data. The active lag distance was set to 40 m (50 % of the

maximum transect length). The lag class distance interval (step size) was selected to

provide the best autocorrelation fit for each individual data set and varied between 2 and

6 m. Anisotropic methods were used as the variation in nutrient concentration was

expected to vary differently between the primary axis of water flow (north-south) and the

61 axis of lateral water flow (east-west). To account for this variation, an offset angle of 90o with an offset tolerance of 22.5o was used for all geostatistical analyses. Interpolation values for a specific location were weighed by distance and the degree of autocorrelation present for that distance. The interpolated isobar maps were created using a uniform x-y grid that generated 188 points at a space interval of 0.6 m and a search neighborhood of

32 nearest points with no limit on geographical distance.

3.4 Results

3.4.1 Hydrologic loading and hydroperiod

Between April 2004 and March 2005there were 8 discrete river pulses into the created wetland that created a hydraulic loading rate of 27 m3 m-2 yr-1 (volume of water

inflow per total area of wetland). From April 2005-March 2006 the inflow was 20 m3 m-2

yr-1 (Table 3.1). The steady state hydrologic year (April 2005-March 2006) had 338

rather that 365 days of inflow due to 4 power outages and general pump failure in the

March 2006. The total inflow into the oxbow wetland was 25% lower in the steady flow

year than in the pulsing year. The experimental design called for the same amount of

water in both years. Most of the discrepancy between the two hydraulic loading rates

occurred due to a loss of pumping ability in March 2006. If this month is disregarded,

then the difference in inflow between the two water years is reduced to 15%.

The important difference between years was likely not the amount of water, but

rather the timing of the water delivery (Fig. 3.3). This difference in delivery is quantified

by the differences in the average monthly Richardson-Baker flashiness index and by the

differences in the mean hydrologic retention time (HRT) between the pulsing and steady-

62 flow years. The index during the pulsing year was substantially greater than the index

during the steady-flow year, 0.96 ± 0.06 and 0.21 ± 0.02 respectively, indicating that the

wetland hydroperiod was indeed much more variable during the pulsing conditions

(Table 3.1). The HRT was different between the two hydroperiods, with the pulsing

hydroperiod condition having a shorter HRT (6 ± 1 days), than the steady-flow

hydroperiod condition, (13 ± 1; Table 3.1). During pulsing hydrologic conditions, 95 %

of the total number of days of flow occurred during the wet season compared to 62%

during non-pulsing conditions. The pulsing flow in 2004-2005 is typical of how this

wetland has historically functioned (Fink and Mitsch, in press).

3.4.2 Nutrient loading and retention rates

The loading rate for nitrate-nitrogen was three times higher during the 2004

pulsing year (32.2 ± 0.2 g-N m-2 yr-1; grams per area within wetland boundary per year) than in the 2005 steady-flow year (10.3 ± 0.1 g-N m-2 yr-1; Table 3.2). The loading rate

was also higher during the pulsing year for total nitrogen (64.5 compared to 17.2 g-N m-2

yr-1) and total phosphorus (6.1 compared to 3.96 g-P m-2 yr-1). The retention rate was

lower for these three parameters during the steady-flow year. Retention rates were 15.4 ±

-2 -1 -2 -1 -2 -1 - 0.2 g-N m yr , 32.3 ± 0.2 g-N m yr , and 4.48 ± 0.03 g-P m yr for NO3 -N, TN, and

TP respectively during the pulsing year and 8.0 ± 0.2 g-N m-2 yr-1, 11.0 ± 0.2 g-N m-2

-1 -2 -1 - yr , and 2.87 ± 0.02 g-P m yr for NO3 , TN, and TP respectively during the steady-

flow year. The only nutrient species that did not have a difference in retention rate was

soluble reactive phosphorus. The loading rates were 0.48 ± 0.02 g-P m-2 yr-1

63 in the pulsing year and 0.14 ± 0.01 g-P m-2 yr-1 in the steady flow year with retention rates of 0.05 ± 0.01 g-P m-2 yr-1 and 0.07 ± 0.2 g-P m-2 yr-1 respectively.

It is not believed that the difference in hydrologic loading rate between the pulsing hydroperiod year and the steady-flow year caused by the pump failure invalidate the results of this study. If the mean inflow concentrations of nitrate, TN, SRP, and TP from March 2004 and 2005 are used as a proxy to calculate likely nutrient loadings for

March 2006 (assuming the target hydraulic loading rate of 200 gallons per minute) it can be seen that the loading rate for the steady-flow hydroperiod year is only expected to increase by 7.5 % for nitrate, 9.2 % for TN, 6.7 % for SRP, and 1.8 % for TP. In the case of each nutrient measured, the difference in loading from pump failure in Marsh is an order of magnitude less than the difference in mean loading rate for the entire year and an order of magnitude less than the difference in nutrient removal rate for either hydroperiod.

3.4.3 Seasonal and spatial nutrient dynamics

There were differences between the spatial dynamics of different nutrient species in pulsing hydroperiod and non-pulsing hydroperiod years (Table 3.3; Fig. 3.4) and between the wet and dry seasons (Figs. 3.5, 3.6). The concentration of nitrate at the wetland inflow (2.87 ± 0.10 mg L-1) was significantly higher (p = 0.05) during pulsing hydroperiod conditions than during steady-flow (2.13 ± 0.07 mg L-1). During the dry season in the steady-flow year there were no difference in nitrate concentration spatially throughout the wetland (p > 0.05; Fig. 3.6). During the pulsing hydroperiod year, there was a slight increase in nitrate concentration in the open water basin during the wet

64 season (Fig 3.5). This did not occur during the steady flow conditions. Under flood pulse conditions, nitrate concentrations in the dry season initially decreased from 1.25 ±

0.10 mg L-1 at the inflow to 0.50 ± 0.05 mg L-1 at the middle of the open water basin before slightly increasing to 0.87 ± 0.10 mg L-1) in the shallow swale that exists between the open water basin and the outflow of the wetland (Fig. 3.6). A similar pattern of increasing nitrate was seen in the wet season of the pulsing year (Fig. 3.5).

It is unclear if the elevated nitrate concentration seen along the west bank of the created oxbow near the outflow is an artifact of the sampling method or if nitrate was produced within the wetland. Because the sampling method was not continuous, it is possible that the grab sampling did not accurately capture each flood pulse as it traveled through the wetland. During most flood-pulses there was an increase in the nitrate concentration at the inflow as nutrient enriched river water flooded into the created oxbow (Fig. 2.6). As water moved through a wetland, some amount of the nitrate can be taken up by plants or transformed by denitrifying bacteria in to N2 gas and emitted into the atmosphere (Mitsch and Gosselink, 2000). The nitrate that was not retained or removed flowed through the wetland as a nutrient pulse whose magnitude dropped as it moved through the wetland. If by chance grab sampling for nutrients occurred near the end of this pulse, it would be possible for the peak of the pulse to be near the outflow, making it appear as though nitrate was being produced by the wetland when in reality some amount was simply moving through untransformed.

Nitrate could also have been produced within the wetland. Soluble organic nitrogen from decaying plants and algae can be mineralized under both aerobic and

+ anaerobic pathways through ammonification to form ammonium ions (NH4 ).

65 Ammonium ions can then be taken up directly by anaerobic organisms, converted to ammonia (NH3) and lost to the atmosphere via volatilization, or it can be immobilized through ion exchange with negatively charged soil particles. Ammonium ions can also be oxidized in the thin oxidized layer that exists at the surface of many wetlands soils by

- Nitrosomonas sp. microorganisms into nitrate (NO2 ) and then by Nitrobacter sp.

- microorganisms into nitrate (NO3 ) in a process called nitrification (Mitsch and

Gosselink, 2000). Nitrification can also occur in the oxidized rhizosphere of plants where adequate oxygen is often available to convert the ammonium nitrogen to nitrate nitrogen (Reddy and Graetz, 1988)

There was a significant difference in the mean total nitrogen concentration at the inflow of the wetland (p = 0.05; Table3.3). During flood-pulse conditions, the mean inflow was 3.04 ± 0.05 mg L-1 and 3.39 ± 0.09 mg L-1 during steady-flow conditions. In

2004, there was no decrease in TN through the emergent marsh portion of the wetland

(Table 2.3) due to an increase in the organic-N (as measured by TKN) concentration (Fig.

3.4). This increase in TN was most pronounced during the dry season (July-October) when there were significant increases during both pulsing and steady-flow hydroperiod conditions (Fig 3.6). In 2004, TN increased from 0.90 ± 0.09 mg L-1 to 3.2 ± 0.09 mg L-1 while in 2005 TN increased from 0.60 ± 0.03 mg L-1 to 3.50 ± 0.30 mg L-1 (Fig 3.4).

There was not a substantial difference in the pattern of the TN dynamics between years and seasons. During both the pulsing and steady flow conditions, after the TN concentration increase through the emergent marsh it then decreased across the open water basin, with a higher concentration band along the more vegetated western shore of the wetland close to the outflow (Figs 3.4, 3.5, 3.6).

66

There was a significantly greater mean concentration (p = 0.05) of SRP at the

wetland inflow during the pulsing hydroperiod (46 ± 4 μg L-1) year than in the steady-

flow (29 ± 1 μg L-1) year (Table 3.3). There was no significant difference in

concentration (p > 0.05) between years, however, at the wetland outflow (Table 2.3).

During the wet season there was a general decrease in SRP under steady flow conditions,

while no spatial differences in concentration were detected during flood pulse conditions

(Fig. 3.5). In the dry season, however, SRP concentration increased across the open water basin during the steady flow conditions whereas it was observed to decrease across that same basin during the pulsing conditions (Fig. 3.6).

There was no significant difference in Total Phosphorus concentration at the inflow of the wetland between the flood-pulse and steady-flow conditions (Table 2.3).

Total phosphorus increased in concentration through the emergent marsh portion of the wetland during the wet season under pulsing conditions (from 143 ± 4 μg L-1 to 203 ± 20

μg L-1), but decreased through this portion of the wetland under all other conditions (Fig.

3.5). While the annual mean TP concentration decreased from 192 ± 16 μg L-1 to 145 ± 5

μg L-1 (18%) during flood-pulse conditions (Table 2.3), this was not the case for all

seasons. During the dry season in the flood-pulse hydroperiod year, the mean TP

concentration did not decrease across the open water portion of the wetland (Fig. 3.6).

The concentration of total suspended solids (18,2 ± 1.8 mg L-1) at the inflow of

the wetland was not significantly different (p > 0.05) between the flood-pulse and the

steady-flow hydroperiod years (Table 2.3). There was, however, a significant (p = 0.05)

67 difference in percent removal (by concentration). In 2004, the outflow TSS concentration

was 8.7 ± 0.2 mg L-1 (52% decrease) while in 2005 it was 13.2 ± 0.8 mg L-1. In both

2004 (250 ± 40 mg L-1) and 2005 (203 ± 20 mg L-1) there was a mean increase in TSS in

the middle of the open water basin (Fig. 3.4). This increase in TSS occurred during

different seasons in the different hydrologic conditions. In the flood-pulse hydroperiod

year (2004) the increase occurred during the dry season (Fig. 3.6) whereas in the steady-

flow year it occurred during the wet season (Fig. 3.5).

3.4.4 Avian use

From April 2004 to March 2005 (flood pulse year), a total of 21 birds species

were observed utilizing the wetland while from April 2005 to March 2006 (steady flow

year), 14 bird species were observed (Table 3.4). A comparison of the number of new

species observed per sampling effort was done (Fig. 3.7). The largest difference in new

species observed between the two hydroperiods occurred during April-May and

September-November. In the spring, the guilds of bird species with the greatest

differences in presence between the pulsing and steady-flow conditions were the geese

and ducks and the songbirds. In the late summer and early autumn the guild of bird

species with the greatest difference in presence between the pulsing and steady-flow

conditions was the shorebirds. During steady-flow conditions there was no exposed

mudflat during the late summer and autumn months. Without this habitat, “peep” type

shorebirds did not utilize the wetland.

68

3.5 Discussion

3.5.1 The Effect of Pulsing

Pulses can be both a subsidy and a stress for an ecosystem. The springtime flood

pulses bring in nutrients (N-P), propagules, colonizing animals, and fresh sediments but the high water can also cause oxygen deprivation in the substrate, impede macrophyte germination, and flood bird nests. This study suggests that a pulsing hydroperiod increases some ecosystem functions. Other studies on pulsing, and other types of disturbances within ecosystems, suggest that some intermediate amount of pulsing may maximize ecosystem function (e.g. Connell, 1978; Mitsch and Ewel, 1979; Odum, 1995;

Townsend, 1996).

The reason for the difference in the loading rates between the pulsing and non- pulsing condition is the difference in the timing and amount of water delivery. This causes a difference in loading rates because there are differences in the concentrations of nitrogen and phosphorus in the river during the different seasons (Figs. 3.5, 3.6). The concentration of nitrate and total phosphorus are typically greater in the spring, following farm field fertilization (Randell et al., 1997; Fink and Mitsch, 2004). This pattern was also observed in this study with the greatest amount of water flowing into the wetland during the pulsing hydroperiod year generally coinciding with elevated concentrations of nitrate and total phosphorus in the Olentangy River. If the maximum delivery of water to the wetland is not timed with the maximum concentration of nutrients in the influent water, then the efficacy of the wetland to treat the nutrient pollution will not be maximized.

69

3.5.2 Comparison to other river diversions

The rates of nitrate-nitrogen and total phosphorus removal are within the ranges

-2 -1 of normal nutrient removal rates in wetlands of 10 - 40 g-NO3-N m yr and 0.5 - 5 g-P

m-2 yr-1 reported by Mitsch et al. (2000). The mean nitrate-nitrogen removal rate of 15.4

± 0.2 g-N m-2 yr-1 in this wetland is at the low end of the range predicted by that paper

while the mean phosphorus retention rate of 32.3 ± 0.2 g-P m-2 yr-1 is at the high end.

Long-term experience with the adjacent experimental wetlands at this same Ohio location

(see Mitsch et al., 2005b) suggest that nitrate-nitrogen removal has increased or remained

steady over a decade while total phosphorus retention has decreased over that time. The

total nitrogen removal rate is also similar to those reported by other studies of river-fed

created wetlands (Table 3.5).

In a whole ecosystem comparison of pulsing and steady flow conditions in created riparian wetlands adjacent to the one described here, Mitsch et al. (2005c) observed

- significant reduction in SRP but no difference in the percent change for NO3 or TP

reduction during non-pulsing conditions. This contrasts with the pattern observed in this

study of no difference in TP percent reduction (by concentration) but more reduction (by

- concentration) in SRP, NO3 , and TN concentrations during pulsing conditions (Table

3.3). This difference may be due to differences in experimental design. In the study

described by Mitsch et al. (2005c) flood pulses were provided artificially by pumping

river water into the wetland during the first week of the month during the wet season

regardless of river flow. Flood pulses were therefore not necessarily synchronized with

70 nutrient pulses in the river. In this parallel study, flood pulses naturally occurred and were therefore more likely to be in sync with elevated nutrient levels in the river.

Studies of river diversion wetlands in the lower region of the Missouri-Ohio-

Mississippi (MOM) watershed have shown similar results. On the Mississippi River in

Louisiana studies of the Caernarvon river diversion showed an 88 to 97 percent reduction

- of NO3 in river water flowing into the receiving wetland basins with a loading rate that ranged from 5.6 to 13.4 g m-2 yr-1 (Lane et al., 1999). Another study during the 1997 opening of the Bonnet Carre Spillway, also on the Mississippi River in LA, showed that

- -2 -1 when river water at a NO3 loading rate of 8.6 g m yr was allowed to flood into the

- LaBanche wetlands and Lake Pontchartrain there was a 92 to 98% reduction of NO3

(Day et al., 1999). A similar study of the same spillway also showed that nutrient concentrations generally decreased, albeit at a lesser rate, with a 28–42% reduction in nitrate, 26–30% in TN, and 50–59% decrease in TP. In the studies compared here, the chemical form of the nitrogen is not an issue as the predominant form of inorganic- nitrogen in Olentangy River water in Ohio and Mississippi River water in Louisiana is

- NO3 with NH3 being only a small fraction (Fink and Mitsch, in press; Day et al., 1999;

Lane et al., 2001; 2002).

3.5.3 Spatial patterns of nitrogen removal in pulsing river diversion wetlands

Total nitrogen increased in the emergent marsh section and subsequently decreased in the open water section of the oxbow wetland in this study. This same pattern was observed in all seasons and during both the pulsing and steady flow hydroperiods (Figs. 3.4, 3.5, 3.6). Because nitrate concentrations decreased through the

71 emergent marsh section of the wetland and measured ammonia concentrations in the

wetland were negligible, this 50% to 100% increase in total nitrogen in the upper reach of

the wetland can be interpreted as an increase in organic-N.

Other river diversion studies have seen increases in mean total Kjeldhal nitrogen

(TKN) concentration in the upper reaches of the wetland area than in the associated river.

Lane et al. (1999) found that TKN increased 33% through the initial portion of an

estuarine wetland in Louisiana before decreasing again through the lower portion. Like

+ this study, NH4 accounted for a small percentage of the TKN (15%), indicating that the

bulk of the TKN concentration is organic-N. Since the same pattern of organic-N

production in the early portion of a nitrogen enriched river diversion wetland followed by

a reduction in organic-N once the NOx-N concentration is reduced was seen in river diversion wetlands at both the top and bottom of the Mississippi watershed, it suggests that this is a pattern that might be generally expected in these types of wetland-river systems.

3.5.4 Management implications

Even with a complete diversion of river floodwaters into the delta, potential nitrate reduction is likely limited to less than 10% to 15% of total nitrate flux in the river

(Day et al., 2000). Thus, if nitrogen flux to the Gulf of Mexico is to be substantially reduced, diversions need to be located in all reaches of the watershed. The creation of diversions for river floodwaters in the upper and lower reaches of the Mississippi-Ohio-

Missouri (MOM) watershed could not only reduce the rate of inland wetland loss but may also reduce nitrogen flux to the Gulf of Mexico (Lane et al. 1999, Perez and Day, 2000).

72 - One of the reasons for the variability in the retention rates of NO3 reported in the

literature is that at high loading rates, removal efficiencies can decrease (Richardson and

Nichols, 1985; Faulkner and Richardson, 1989; Boustany et al., 1997; Spieles and

Mitsch, 2000). For example, Spieles and Mitsch (2000) found in the upper reaches of the

- MOM watershed only a 37 to 40% reduction in NO3 in wetlands receiving Olentangy

River water at loading rates of 4.6 to 4.7 kg ha-2 day-1 (equivalent to 168 to 172 g m-2

yr-1). For another example from the lower reaches of the MOM watershed, in 1997, the

Atchafalaya River -wetland complex had a loading rate of 66 to 136 g m-2 yr-1,

- with a 41 to 47% decrease in NO3 (Lane et al., 2002).

The way that water flows through a system is also very important. In many wetlands, flowing water forms small channels and the area of actual water-soil contact is much lower than the total areal receiving area, which limits the exposure of nitrate to denitrifying bacteria.. For example, Blahnik and Day (2000) showed that for a treatment wetland in Louisiana, that 60% of the surface water only contacted 10-12% of the total area of the wetland. Since the way to achieve maximum efficiency of nitrogen removal is for diverted water be spread over as much of the wetland as possible, care must be given during design to minimize the possibility of channel formation with the subsequent

“short-circuiting” of the wetland.

3.5.5 Avian effects

Flood pulsing also had an impact on the use of the wetland by different species of birds. Traditionally, if wetlands in the Midwest have been managed for birds, they have been managed for waterfowl (Fredrickson and Taylor, 1982; Taft et al., 2002). The loss

73 of drawdown conditions that occurred in this study when a pulsing hydroperiod was replaced with a steady-flow had two noticeable effects on avian use. First, herons were observed feeding a fewer number of times in the wetland in the late summer and early fall. During this time in the year with a pulsing hydroperiod, the open water area of the created oxbow was one-half to one-third its maximum size in terms of areal coverage.

Perhaps fish were concentrated into a smaller pool of water, creating a more appealing food density for herons and egrets. Second, the wetland did not have any exposed mudflat in the late summer and early fall in the year in which pulses were removed. The lack of mudflat corresponded with fewer observations of shorebird species. This is important as this time period of natural drawdown and mudflat exposure matches the time of shorebird migration (Poole et al.; 1992). River diversion wetlands that allow natural seasonal pulses may provide critical feeding habitat for these migrating shorebirds.

3.6 Conclusions

Delivering water to wetland with natural flood pulses is likely to synchronize nutrient and water pulses in most years. Our study suggests that this may maximize the nutrient removal potential of the wetland for the watershed. A wetland with a riverine flood-pulse inflow may experience isolation from its supplying river in the late summer during the dry season. This will likely have a minimal impact on nutrient removal within the watershed as a whole because the concentrations of nutrients in temperate rivers are often low at this time of year. Furthermore, the hydrologic isolation from the river allows for drawdown conditions and mudflat exposure, which allows for migrating shorebirds to utilize the wetland and for plants to grow in more areas of the wetland in the summer.

74 Our research suggests that by removing the pulsing hydroperiod, the wetland loses temporal habitat variability. Specifically the wetland loses the drawdown portion of its annual hydroperiod and the wetland edge becomes static. Restoring a flood-pulse hydroperiod to riparian wetland in the Mississippi watershed may increase the rate of nutrient removal and the amount of avian use within river diversion wetlands.

3.6 Acknowledgements

Support provided by U.S. Department of Agriculture NRI CSREES Award 2003-

35102-13518 and a Payne grant from the Ohio Agricultural Research and Development

Center of The Ohio State University. We also thank Maria Hernandez for coordinating the laboratory analysis of nutrient concentrations.

75 3.7 Literature cited

American Public Health Association. 1998. Standard Methods for the Analysis of Wastewater, 19th ed. APHA, Washington DC.

Baker, David B., R. Peter Richards, Timothy T. Loftus, Jack W. Kramer. 2004. A new flashiness: Characteristics and applications to Midwestern rivers and streams. Journal of the American Water Resources Association 40:503-522.

Bedford, B.L. 1996. The need to define hydrologic equivalence at the landscape scale for freshwater wetland mitigation. Ecological Applications 6:57-68.

Blahnik, T. and J. Day. 2000. The effects of varied hydraulic and nutrient loading rates on water quality and hydrologic distributions in a natural forested treatment wetland. Wetlands 20:48-61.

Boustany, R.G., C.R. Croizer, J.M. Rybczyk, R.R. Twilley. 1997. Denitrification in a south Louisiana wetland forest receiving treated sewage effluent. Wetlands Ecology and Management 4:273-283.

Connell, J.H. 1978. Diversity in tropical rainforest and coral reefs. Science 199:1302- 1310.

Day, J. W., Jr., T. J. Butler, and W. G. Conner. 1977. Productivity and nutrient export studies in a cypress swamp and lake system in Louisiana. pp. 255-269. In: M. Wiley, (ed.) Estuarine Processes, Vol. II. Academic Press, New York.

Day, J.W., R.R. Lane, R.F. Mach, C.G. Brantley, M.C. Daigle. 1999. Water chemistry dynamics in Lake Pontchartrain, Louisiana, during the 1997 opening of the Bonnet Carre Spillway. Proceedings of Recent Research in Coastal Louisiana. Lafayette, LA.

Day, J.W., L.D. Britsch, S. Hawes, G. Shaffer, D.J. Reed, and D. Cahoon, 2000. Pattern and process of land loss in the Mississippi Delta: a spatial and temporal analysis of wetland habitat change. Estuaries 23:425-438.

Fink, D.F. and W.J. Mitsch. 2004. Seasonal and storm event nutrient removal by a created wetland in an agricultural watershed. Ecological Engineering 23:313-325.

Fink, D.F. and W.J. Mitsch. in press. Hydrology and nutrient biogeochemistry in a created river diversion oxbow wetland. Ecological Engineering

Faulkner, S.P., C.J. Richardson. 1989. Physical and chemical characteristics of freshwater wetland soils. pp. 41-72. In: Hammer, D.A. (ed.), Constructed Wetlands for Wetland . Lewis Publishers.

76 Fernandez, J.M., M.A.E. Selma, F.R. Aymerich, M.T.P. Saez, M.F.C Fructuoso. 2005. Aquatic birds as indicators of trophic changes and ecosystem deterioration in the Mar Menor lagoon (SE Spain). Hydrobiologia 550:221-235.

Fredrickson, L.H. and T.S. Taylor. 1982. Management of Seasonally Flooded Impoundments for Wildlife. Resource Publication 148. US Fish and Wildlife Service, Washington, DC.

Galat D.L., L.H. Frederickson, D.D. Humburg, K.J. Bataille, J.R. Bodie, J. Dohrenwend. 1998. Flooding to restore connectivity of regulated, large-river wetlands. (Lower Missouri River). BioScience 48:721–733.

Gibbs, J.P., J.R. Longcore, D.G. McAuley, and J.K. Ringelman. 1991. Use of wetland habitats by selected nongame waterbirds in Maine. U.S. Fish and Wildlife Service Fish Wildlife Resources 9. 57 p.

Harter, S.K. and W.J. Mitsch. 2003. Patterns of short-term sedimentation in a freshwater created marsh. Journal of Environmental Quality 32:325-334.

Hein, T., G. Heiler, D. Pennetzdorfer, P. Riedler, M. Schageri, F. Schiemer. 1999. The Danube Restoration Project: functional aspects and plantonic productivity in the floodplain system. Regulated Rivers 15:259-279.

Junk, W. J. 1999. The flood pulse concept of large rivers: Learning from the tropics. Archiv für Hydrobiologie 115: 261–280.

Lachet Instruments. 2000. Methods Manuel. Lachet Instruments, Milwaukee, WI, USA.

Lane, R.R., J.W. Day, B. Thibodeaux. 1999. Water quality analysis of a freshwater diversion at Caernarvon, Louisiana. Estuaries 22:327-336.

Lane R.L., J.W. Day, G.P. Kemp, D.K. Demcheck. 2001. The 1994 experimental opening of the Bonnet Carre Spillway to divert Mississippi River water into Lake Pontchartrain, Louisiana. Ecological Engineering 17, 411–422

Lane, R.R., J.W. Day, G.P. Kemp, B. Marx, E. Reyes. 2002. Seasonal and spatial water quality changes in the outfall plume of the Atchafalaya River, Louisiana, USA. Estuaries 25, 30-42.

Lane, R.L., J.W. Day, D. Justica, E. Reyes, B. Marx, J.N. Daya, E. Hyfield. 2004. Changes in stoichiometric Si, N and P ratios of Mississippi River water diverted through coastal wetlands to the Gulf of Mexico. Estuarine, Coastal and Shelf Science 60:1-10

77 McDougal, R.L., L.G. Goldsborough, B.J. Hann. 1997. Responses of a prairie wetland to press and pulse additions of inorganic nitrogen and phosphorus: Production by planktonic and benthic algae. Archiv für Hydrobiologie 140:145-167.

Megonigal, J. P., W. H. Conner, S. Kroeger, R. R. Sharitz. 1997. Aboveground production in southeastern floodplain forests: A test of the subsidy-stress hypothesis. Ecology 78:370-384.

Mitsch, W.J. and K.C. Ewel. 1979. Comparative biomass and growth of cypress in Florida wetlands. American Midland Naturalist 101:417-426.

Mitsch, W.J., C.L. Dorge, J.W. Wiemhoff. 1979. Ecosystem dynamics and a phosphorus budget of an alluvial cypress swamp in southern Illinois. Ecology 60:1116-1124.

Mitsch, W.J. and W.G. Rust. 1984. Tree growth responses to flooding in a bottomland forest in northeastern Illinois. Forest Science 30:499-510.

Mitsch, W.J., J.W. Day, J.W. Gilliam, P.M Groffman, D.L Hey, G.W. Randell, N. Wang. 1999. Reducing nutrient loads, especially nitrate-nitrogen, to surcace water, groundwater, and the Gulf of Mexico. Topic 5 Report for the Integrated Assessment on Hypoxia in the Gulf of Mexico. NOAA Coastal Ocean Program Decision Analysis Series No. 19. NOAA Coastal Ocean Program, Silver Spring, MD, 111 pp.

Mitsch, W.J and J.G. Gosselink. 2000. Wetlands, 3rd ed. John Wiley and Sons, NY.

Mitsch, W.J., A.J. Horne, R.W. Nairn. 2000. Nitrogen and phosphorus retention in wetlands —Ecological approaches to solving excess nutrient problems. Ecological Engineering 14, 1-7.

Mitsch W.J., J.W. Day, J.W. Gilliam, P.M Groffman, D.L Hey, G.W. Randell, N. Wang. 2001. Reducing nitrogen loading to the Gulf of Mexico from the Mississippi River Basin: Strategies to counter a persistent ecological problem. BioScience 51:373-388.

Mitsch, W.J., N. Wang, L. Zhang, R. Deal, X. Wu, A. Zuwerink. 2005a. Using ecological indicators in a whole-ecosystem wetland experiment. In: Jorgensen, S.E., F.L. Xu, R. Costanza (Eds.), Handbook of Ecological Indicators for Assessment of Ecosystem Health, CRC Press, Boca Raton, FL, PP.211-235.

Mitsch, W.J., J.W. Day, Jr., L. Zhang, R. Lane. 2005b. Nitrate-nitrogen retention by wetlands in the Mississippi River Basin. Ecological Engineering 24:267-278.

Mitsch, W.J., L. Zhang, C.J. Anderson, A.E. Altor, M.E. Hernandez. 2005c. Creating riverine wetlands: Ecological succession, nutrient retention, and pulsing effects. Ecological Engineering 25:521-527.

78 Mitsch, W.J. and J.W. Day. 2006. Restoration of wetlands in the Mississippi-Ohio- Missouri (MOM) River Basin: Experience and needed research. Ecological Engineering 26:55-69.

Nixon, S. W., J.W. Ammerman, L.P. Atkinson, V.M. Berounsky, G. Billen, W.C. Boicourt, W.R. Boynton, T.M. Church, D.M. Ditoro, R. Elmgren, J.H. Garber, A.E. Giblin, R.A. Jahnke, N.J.P. Owens, M.E.Q. Pilson, S.P. Seitzinger. 1996. The fate of nitrogen and phosphorus at the land-sea margin of the North Atlantic Ocean. Biogeochemistry 35:141-180.

Odum W.E., E.P. Odum, H.T. Odum. 1995. Nature’s pulsing paradigm. Estuaries 18:547–555.

Perez, B. C. and J. W. Day. .2000. Influence of Atchfalaya River discharge and winter frontal passage and flux in Four League Bay, Louisiana. Estuarine. Coastal and Shelf Science 50: 271-290.

Poole, A.F., P.R. Stettenheim, F. . 1992. The Birds of North America: Life Histories for the 21st century. (eds) Alan F. Poole, Peter Stettenheim, and Frank Gill. American Ornithologists' Union; Washington, D.C.

Rabalais, N. N., W. J. Wiseman, R. E. Turner, B. K. Sengupta, and Q. Dortch. 1996. Nutrient changes in the Mississippi River and system responses on the adjacent continental shelf. Estuaries 19:386-407.

Rabalais, N. N., R. E. Turner, W. J. Wiseman, Q. Dortch. 1998. Consequences of the 1993 Mississippi River flood in the Gulf of Mexico. Regulated Rivers 14:161-177.

Rabalais, N.N., R.E. Turner, D. Justic, Q. Dortch, W.J. Wiseman. 1999. Topic 1 Report for the Integrated Assessment on Hypoxia in the Gulf of Mexico. NOAA Coastal Ocean Program Decision Analysis NO. 15; NOAA Coastal Ocean Program, Silver Spring, MD,. 167PP.

Rabalais, N.N., R.E. Turner, D, Scavia. 2002. Beyond science and into policy: Gulf of mexico hypoxia and the Mississippi River. BioScience 52:129-142.

Randell, G.W., D.R. Huggins, M.P. Russelle, D.J. Fuchs, W.W. Nelson, J.L Anderson. 1997. Nitrate losses through subsurface tile drainage in CRP, alfalfa and row crop systems. Journal of Environmental Quality 26:1240-1247.

Reddy, K.R. and D.A. Graetz. 1988. Carbon and nitrogen dynamics in wetland soils. Pp. 307-318. In: D.D. Hook, W.H. McKee, Jr., H.K. Smith, J. Gregory, V.G. Burrel, M.R. DeVoe, R.E. Sojka, S. Gilbert, R.Banks, L.G. Stolzy, C. Brooks, T.D. Mathews, and T.H Shear (eds.) The Ecology and Management of Wetland, Vol. 1: The Ecology of Wetlands. Timber Press, Portland, OR.

79 Reyes, E, J.F. Martin, J.W. Day, G.P. Kemp, H. Mashriqui. 2004. River forcing at work: ecological modeling of prograding and regressive deltas. Wetlands Ecology and Management 12:103-114.

Richardson, C.J., D.S. Nichols. 1985. Ecological analysis of wastewater management criteria in wetland ecosystems, Ecological considerations In: E.R.K. Paul J. Godfrey, Sheila Pelczarski (ed.), Wetlands Treatment of Municipal Wastewaters, Van Nostrand Reinhold Company, New York, PP. 351-391.

Rutschke, E., 1987. Waterfowl as bio-indicators. In: Diamond, A.W., F.L. Filion (eds), The value of birds. ICBP Technical Publication No. 6. 167–172.

Spieles, D.J. and W.J. Mitsch. 2000. The effects of season and hydrologic and chemical loading on nitrate retention in constructed wetlands: a comparison of low and high nutrient riverine systems. Ecological Engineering 14:77-91.

Steen, D.A., J.P. Gibbs, T.A. Timmermans. 2006. Assessing the sensitivity of bird communities to hydrologic change in the eastern Great Lakes region. Wetlands 26:605-611.

Taft, O.W., M.A. Colwell, C.R. Isola, R.J. Safran. 2002. Waterbird responses to experimental drawdown: implications for the multispecies management of wetland mosaics. Journal of Applied Ecology 39:987-1001.

Tockner , K., F. Malard, J.V. Ward. 2000. An extension of the flood pulse concept. Hydrologic Processes 14:2861-2883.

Townsend, C.R. 1996. Concepts in river ecology: pattern and process in the catchment hierarchy, Archiv Fur Hydrobiologie, Supplemental.

Turner, R. E., N.N. Rabalais, E.M. Swenson, V. Kasprzak, I. Romaire, T. Marine. 2005. Summer hypoxia in the northern Gulf of Mexico and its prediction from 1978 to 1995. Environmental Research 59:65-77.

U.S. Environmental Protection Agency. 1983. Handbook for Methods in Water and Wastewater Analysis. U.S. Environemntal Protection Agency, Cincinnati, OH.

Vannote, R.L., G.W. Minshall, K.W. Cummins, J.R. Sedell, C.E. Cushing. 1980. The river continuum concept. Canadian Journal of Fisheries and Aquatic Science. 37:130-137.

Wilcox, D. A. 1993. Effects of water-level regulation on wetlands of the Great Lakes. Great Lakes Wetlands 4:1-11.

Wilcox, D.A. and T.H. Whillans. 1999. Techniques for restoration of disturbed coastal wetlands of the Great Lakes. Wetlands 19:835-857.

80

Figure 3.1 Site map of the Wilma H. Schiermeier Olentangy River Wetland Research

Park on The Ohio State University campus. The 2.8 ha created oxbow wetland is between the experimental wetlands and the bottomland hardwood forest. Grab-sample locations are marked with a white circle. The area to the north of the “created oxbow” label is predominantly emergent marsh and the area to the south of the label is predominantly open water.

81

82

Figure 3.2 Inflow (top) and outflow (bottom) control structures in the created oxbow wetland. The Red Field TideflexTM check valve, top, opens via water pressure when the river elevation is higher than the wetland and closes when the river elevation is lower than the wetland water level, and water pressure is removed. Water then flows back to the Olentangy River though an outflow control weir, bottom.

83

84

Figure 3.3 Water inflow rate for the created oxbow wetland between April 2004 and

March 2006. April 2004 through March 2005 was a pulsing hydroperiod year while April

2005 through March 2006 was steady-flow hydroperiod year when flood-pulses were removed from the wetland.

85

86

- Figure 3.4 Kriging diagrams of the nitrate + nitrate (NO3 -N), total nitrogen (TN), soluble reactive phosphorus (SPR), total phosphorus (TP), and total suspended solids (TSS) in the created oxbow wetland showing annual mean concentrations for 2004 and 2005.

Inflow of river water is on the left of each diagram; outflow is on right. Sampling locations are shown in Fig. 3.1.

87 0 200 N 0 60 feet meters

88

- Figure 3.5 Kriging diagrams of the nitrate + nitrate (NO3 -N), total nitrogen (TN), soluble reactive phosphorus (SPR), total phosphorus (TP), and total suspended solids (TSS) in the created oxbow wetland during the wet season in 2004 and 2005. Inflow of river water is on the left of each diagram; outflow is on right. Sampling locations are shown in

Fig. 3.1.

89

0 200 N 0 60 feet meters

90

- Figure 3.6 Kriging diagrams of the nitrate + nitrate (NO3 -N), total nitrogen (TN), soluble reactive phosphorus (SPR), total phosphorus (TP), and total suspended solids (TSS) in the created oxbow wetland during the dry season in 2004 and 2005. Inflow of river water is on the left of each diagram; outflow is on right. Sampling locations are shown on Fig.

3.1.

91

0 200 N 0 60 feet meters

92

Figure 3.7 Number of new avian species-observed per sampling effort in the created oxbow wetland with a pulsing hydroperiod (April 2004 – March 2005) and steady-flow hydroperiod (April 2005 – March 2006).

93

30 April 2004 - March 2005 pulsing April 2005 - March 2006 steady flow 25

y = 6.9727Ln(x) + 0.1238 R2 = 0.9702 20

15 y = 3.7884Ln(x) + 2.2894 94 R2 = 0.859 Number of species 10

5

0 Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar

Month Sampled

94

Pulsing Steady-flow 2004-2005 2005-2006 Wet Season Days of flow 165 148 Inflow m 19 11 Outflow m 17 11 HRT d 6 ± 1 14 ± 1 Dry Season Days of flow 7 90 Inflow m 8 9 Outflow m 9 9 HRT d 4 ± 1 7 ± 1 Annual Number of Pulses 8 0 Total Days of flow 172 338 Inflow m 27 20 Outflow m 26 20 HRT d 6 ± 1 12 ± 1 R-B index 0.96 ± 0.06 0.21 ± 0.02

Table 3.1 Inflow and outflow of the oxbow wetland in April 2004 – March 2006.

Hydraulic retention time (HRT) was calculated only for days in which there was no inflow or outflow. The Richardson-Baker flashiness index (R-B index) for the created oxbow wetland was calculated on a monthly basis for pulsing and steady-flow hydroperiods. Wet season is defined as November through June; dry season is defined as

July through October.

95

- NO3 -N TN SRP TP

2004 2005 2004 2005 2004 2005 2004 2005

Yearly Mean

Loading Rate g-X m-2 yr-1 32.2±0.2 10.3±0.1 64.5±0.4 17.2±0.1 0.48±0.00 0.14±0.01 6.10±0.04 3.96±0.01

Export Rate g-X m-2 yr-1 16.8±0.2 2.3±0.1 32.2±0.3 6.2±0.1 0.43±0.01 0.06±0.01 1.62±0.01 1.09±0.01

Removal

Rate g-X m-2 yr-1 15.4±0.2 8.0±0.2 32.3±0.2 11.0±0.2 0.05±0.01 0.07±0.2 4.48±0.03 2.87±0.02

Percent (by mass) 48±3 77±6 50±4 64±6 10±1.0 57±5 73±8.0 13±7

96 Unit ‘X’ is N or P, as appropriate

- Table 3.2 Annual mean loading and retention of nitrate-nitrite (NO3 -N), total nitrogen (TN), soluble

reactive phosphorus (SRP), and total phosphorus (TP) during pulsing hydroperiod (2004) and steady-

flow (2005) years in the river diversion oxbow (ave ± std. error). Rates are in g-N m-2 yr-1 or

g-P m-2 yr-1 as is appropriate. Rates and the mass retention during the pulsing year are calculated

according to the number of actual days of flow; note that the duration of the inflow and outflow are

different.

96

Parameter Inflow Mid-point Outflow % Removal

Steady Pulsing Steady flow Pulsing Steady flow Pulsing Steady flow Pulse flow SRP (μg-P L-1) 46.5±4 (141)b 29.0±1 (33)b 22±9 (22)c 15±1 (24)c 23.2±1.3 (184)c 18±1 (31)c 50.1 39.2b

TP (μg-P L-1) 192±16 (81) 170±3 (36) 110±16 (14)c 126±3 (24)c 145±3 (108)b 139±5 (32)b 24.5 18.3

- - NO3 + NO2 2.87±0.10 (141)b 2.13±0.07 (21)b 1.76±0.21 (21)bc 0.97±0.06 (21)bc 1.52±0.03 (199)b 0.76±0.03 (20)b 47.0b 64.2b (mg-N L-1)

TN (mg-N L-1) 3.04±0.05 (77)b 3.39±0.09 (21)b 3.17±0.35 (17)c 2.53±0.11 (20)bc 2.31±0.02 (102)c 1.94±0.08 (20)c 24.0 b 42.7b

TSS (mg L-1) 17.3±2.0 (166) 18.2 ±1.8 (36) 14.3±1.3 (23)b 9.8±0.3 (23)bc 13.2±0.8 (227)bc 8.7±0.2 (35)b 23.7 52.1b

97 bSignificant difference between pulsing and steady flow (p = 0.05)

cSignificant difference from upstream location (p = 0.05)

Table 3.3 Nutrient and suspended sediment concentrations (ave ± std error (# samples) in the created oxbow wetland at the

Olentangy River Wetland Research Park during pulsing (April 2003 – March 2005) and non-pulsing (April 2005 – March

2006) conditions when flooded river water is flowing through the wetland (SRP = soluble reactive phosphorus, TP = total

- - phosphorus, NO3 + NO2 = nitrate and nitrite, TN = total nitrogen, and TSS = total suspended solids).

97

Pulsing Non-pulsing April 2004 - April 2005 - Bird species March 2005 March 2006 Heronlike birds Great egret Ardea alba X X Great blue heron Ardea herodias X X American bittern Botaurus lentiginosis X X Green heron Butorides virescens X X Geese and Ducks duck Aix sponsa X X Northern shoveler Anus clypeata X Blue wing teal Anus discors X X Mallard Anus platyrhynchos X X Canada goose Branta canadensis X X Ruddy duck Oxyura jamaicensis X Shorebirds Least sandpiper Calidris minutilla X Killdeer Charadrius vociferus X X Solitary sandpiper Tringa solitaria X Greater yellowlegs Tringa melanoleuca X Swallows and Kingfishers Kingfisher Ceryle alcyron X X Barn swallow Hirundo rustica X X Tree swallow Tachycinetea bicolor X X Songbirds American goldfinch Carduelis tristis X X Sedge wren Cistothorus platensis X Song sparrow Melospiza melodia X X Louisiana waterthrush Seiurus motacilla X Total species observed 21 14

Table 3.4 Bird species observed utilizing the created oxbow wetland with a pulsing hydroperiod, April 2004 – March 2005, and with a steady-flow hydroperiod, April 2005 – March 2006.

98

Annual removal Type of hydroperiod - -2 -2 Study Site g NO3 -N m g P m Reference Sustainable retention rates 10 – 40 0.5 – 5 Pulsing and Steady-flow Mitsch et al., 2000 for non-treatment wetlands Riparian oxbow, Columbus, OH 15 4.5 Pulsing Fink and Mitsch in press 8 2.9 Steady-flow Fink and Mitsch this study River-fed created wetland, 11-38 1.4-2.9 Pulsing Phipps and Crumpton, 1994 Lake County, IL Mitsch et al., 1995 River-fed created wetland, 58-66 5.2-5.6 Pulsing Spieles and Mitsch, 2000 Columbus, OH Nairn and Mitsch, 2000 99 Mississippi River diversion, 5.6-13.4 0.41-0.92 Pulsing Lane et al., 1999 Caernarvon, LA Day et al., 1999 Mitsch et al., 2005a Atchafalaya Delta, 31-54 Pulsing Lane et al., 2001 LA Mitsch et al., 2005a

Table 3.5 Annual and per pulse removal of nitrate (as N) and total phosphorus in selected created river wetlands.

99

CHAPTER 4

THE EFFECT OF REMOVING AND REINTRODUCING HYDROLOGIC PULSES

ON PRODUCTIVITY OF A RIVER-DIVERSION WETLAND

4.1 Abstract

A whole-ecosystem study was done to examine the effects of removing and

restoring flood pulsing on the net primary productivity of emergent macrophytes in a

created riparian oxbow wetland at the Schiermeier Olentangy River Wetland Research

Park at The Ohio State University in Columbus, Ohio USA. The wetland was received 8

natural river pulses from April 2004 – March 2005, a steady flow of artificially pumped

river water from April 2005 – March 2006, and 6 natural river pulses from April 2006 –

September 2006. The wetland was significantly (α = 0.05) more productive during pulsing hydroperiod conditions. Wetland macrophyte productivity decreased from 7560 kg yr-1`to 6544 kg yr-1and total areal coverage decreased from 32.3% to 25.7% after flood

pulsing was removed. When flood pulsing was restored in the third year of the study, the

total net macrophyte productivity increased to 9916 kg yr-1 and areal coverage returned to

32.1%. There was a positive correlation between the R-B flashiness index of the

wetlands hydrology and productivity (r2 = 0.75). It is likely that a pulsing hydroperiod

should enable mitigation wetlands to reach target macropyte coverage goals more easily.

100 4.2 Introduction

Flood pulses affect emergent macrophyte primary productivity by changing nutrient influx (Brown, 1981), water depth (Waters and Shay et al., 1992; Newman et al.,

1998; Kellogg et al., 2003), and the frequency and duration of inundation (Giovanni and

Da Motta Marques, 1998; Newman et al., 1998; Tanner, 1999; Casanova and Brock,

2000). Ecosystems that receive pulses of energy inputs (e.g. tides, river, floods, pulses of runoff, upwelling) in addition to solar energy are generally more productive than those that do not receive pulses (Mitsch and Ewel, 1979; Nixon, 1988; Odum et al., 1995).

Henry et al. (2002) showed that vegetation composition in a restored river diversion wetland exhibited changes related to hydrologic variations and that the wetland colonized rapidly as a result of plant propagules remaining in the sediments or drifting in as a result of flood pulses. Prolonged inundation from steady-flow hydroperiod also increases the potential for anoxic conditions in the root zone of emergent macrophytes, slowing their growth (van der Valk and Davis, 1978); this suggests that periodic drawdowns may enhance wetland plant growth. Flood pulses from rivers also often contain elevated loadings of nutrients (Fink and Mitsch, in press), which may provide a fertilizing effect for plant growth (Spink et al., 1998; Gathumbi et al., 2005; Gusewell et al., 2003).

Increased nutrient availability may also occur as phosphorus sorbed to the surface of wetland sediments may be released during anaerobic conditions and by the increase in nutrient mineralization caused by fluctuations in wet-dry soils conditions associated with a pulsing hydroperiod (Olila et al., 1997). Day et al. (1977; 1995;2000), Mitsch and Ewel

(1979), Mitsch et al. (1979), Megonigal et al. (1997), McDougal et al. (1997), Galat et al.

(1998), Reyes et al. (2004), Lane et al. (2004) and Mitsch et al. (2005c) describe riverine

101 hydrologic pulses as having both positive and negative effects on overall biotic function of wetland ecosystems. It is important to understand these effects because pulsing water regimes are characteristic of riparian wetlands adjacent to flashy streams and rivers in the

Midwest United States (Baker et al., 2004). The temporary nature of river-fed flood pulses causes the water level in receiving wetlands to both increase and decrease rapidly, increasing the opportunity for nutrient mineralization and decreasing the opportunity of soil anoxia damaging to macrophyte growth (Mitsch and Rust, 1984). While the effects of flood-pulsing on productivity has been demonstrated for forested wetland ecosystems

(Mitsch et al., 1991; Brown, 1981) and for planktonic communities (Cronk and Mitsch,

1994; Hein et al., 1999; Tuttle and Mitsch, in review) the overall effects of pulses on macrophyte productivity in riparian river diversion marsh wetland ecosystems is not clear, nor has it been clearly demonstrated in ecosystem studies (Mitsch and Day, 2006).

This may be because herbaceous wetland vegetation is sensitive to short-term fluctuations in different flood conditions (e.g. seasonal timing, duration, depth, etc), which can make determining the specific effects of pulsing difficult.

4.2.1 Goals and Objectives

This paper has a goal of demonstrating the effects of removing and then reintroducing natural nutrient-laden flood pulses of river water on the vegetation communities in created riparian diversion wetlands on a fourth-order river in central

Ohio, USA. The response of the vegetation in a created oxbow wetland was determined through measurements of macrophyte productivity and community diversity. The following specific objectives fit to this goal.

102

1. Investigate the effect of pulsing vs. non-pulsing hydrology on community

diversity of hydrophytic vegetation in a riparian wetland;

2. Determine the effect on total macrophyte productivity in the riparian wetland

when the original pulsing hydrology of the wetland is removed and replaced by a

steady flow of water;

3. Determine if there is an effect on wetland macrophyte productivity when pulsing

hydrological conditions are restored to the wetland.

4.3 Methods

4.3.1 Site Description

The wetland in this study is a created river diversion wetland located at the Wilma

H. Schiermeier Olentangy River Wetland Research Park at The Ohio State University in

Columbus, Ohio, USA (latitude 40.021°N, longitude 83.017°E; Fig. 4.1). A river diversion wetland is a wetland on the adjacent floodplain or behind artificial levees that receive water by pumping or flood flows from the main channel of a river (Mitsch and

Day, 2006). From its creation in 1996 until 2004, the 3-ha created riparian wetland

(referred to here as a created oxbow) received, on average, 7 to 8 natural flood pulses per year from the Olentangy River. Inflow from 1998-2004 averaged 20 ± 4 m yr-1. The

Olentangy River provides frequent short (5-6 days of inflow) flood pulses into the created oxbow, which typically result in 9-12 days of outflow from the wetland. Water flows into the northern tip of the wetland through a Red Field TideflexTM check valve when the

103 river elevation is higher than the wetland; the valve closes when the river elevation is lower than the wetland water level and water then flows back to the Olentangy River though an outflow control weir (Fink and Mitsch, in press).

4.3.2 Hydroperiod

The 3-ha created oxbow was monitored during one year of natural pulsing followed by a second year when flood pulsing was removed and inflow was maintained as steady-flow, followed by a third year when natural flood pulsing was restored

(Fig. 4.2). The third year of flood pulsing extended through the peak biomass of the growing season when productivity measurements were made. Pulsing hydroperiod conditions during the first year and third year (April 2004 to March 2005 and March

2006 to September 2006) were created entirely by natural flooding of the river itself.

Steady-flow conditions during the second year (April 2005 – March 2006) were created and controlled by a large submersed bypass-pump on the river intake that replaced flood pulses with artificial steady-flow conditions. We attempted to provide a similar volume of water to the wetland in the non-pulsing hydroperiod year with the pump as was provided naturally by the river during the first pulsing year (Chapter 3).

Flows into and out of the wetland were measured using a Swofer 2100 current meter for water velocity and an ISCO 730 bubbler module to estimate stage and cross- sectional area. A simple mathematical model was developed to describe the inflow based up upon the relative elevation of the river and the oxbow water surface. Outflow was

104 estimated from the stage of water within the wetland and the shape of the outflow weir

(USBR, 1997). A daily hydrologic budget was then developed from daily readings of river elevation and wetland stage (Fink and Mitsch, in press).

4.3.3 Macrophyte productivity and diversity

The wetland has two significant vegetative zones. The northern (closest to the inflow) half is an emergent marsh and the southern half (closest to the outflow) is an open water basin. The lack of vegetation in the southern half is primarily due to high water conditions during spring, which prevents the germination of emergent aquatic plants except for littoral zones. Vegetation surveys and peak biomass surveys were conducted in August 2004, 2005, and 2006. Sampling transects for vegetation surveys with a total length of 963 m were conducted throughout the entire wetland basin, covering wetted, transitional, and near upland zones. For each species observed, relative abundance was estimated as present (0-5%), common (5-50%), or abundant (50-100%). Indicator status was determined using the Region I (Northeast) National Wetland Indicator List (Reed,

1998). Species not found on this list were recorded as non-listed (NL). Net aboveground primary productivity (NAPP) was estimated by determining peak biomass along six 15-

87 m transects running shore-to-shore along the narrow axis of the wetland that passed through different zones of the wetland. Three separate 1-m2 plots (18 m2 total) were selected randomly within the areas supporting vegetation along each transect. In each plot the above-ground biomass was harvested by cutting the plant at the base as close to the substrate as possible, separated by species, and weighed. Subsamples were dried at

105oC to calculate a wet-dry ratio. Color aerial photographs taken by the Ohio

Department of Transportation on or about the same time as the biomass harvesting were

105 used to estimate the extent of the different macrophyte communities in the wetland. Total

net primary productivity for the wetland was estimated as the product of plot biomass

estimates and the areal extent of macrophyte communities.

Spatial community diversity was quantified by calculating a macrophyte

community diversity index (CDI) (Mitsch et al., 2005a). The index is expressed as:

CDI = Σ (Ciln(Ci)) (2)

Where Ci is the present cover of community “i” and N is the number of macrophyte

communities. Only communities of plants that could be identified from the aerial

photographs were used in this calculation. This index includes evenness of plant cover as

well as the number of plant communities.

4.3.4 Statistical Methods

Direct comparisons of single variables between the water years of the study were

done using homoscedastic, two-tailed Student’s T-Tests for equal means. All tests were

conducted at a 95% confidence interval (α = 0.05).

4.4 Results

4.4.1 Hydrologic loading and hydroperiod

In 2004 there were 8 discrete river pulses into the created wetland for a hydraulic

loading rate of 27 m3 m-2 yr-1 (cubic meters of inflow per square meter of wetland per year; Fig. 4.2). From April 2005-March 2006 the inflow was 20 m3 m-2 yr-1 (Table 4.1).

106 From April 2006-September 2006 there were 6 discrete pulses into the created wetland

for a 7-month hydraulic loading of 13 m3 m-2. The steady-state hydrologic period (April

2005-March 2006) had 338 days of inflow due to 4 power outages and general pump

failure in the March 2006. The inflow rate into the oxbow wetland was slightly less

(11 %) in the steady-flow year than in the pulsing years.

4.4.2 Macrophyte communities

Overall there were 6 different vegetation communities identified from ground

surveys and aerial photography in each year of the study, 5 of which were herbaceous

macrophyte communities and one of which was woody (Fig. 4.3). The macrophyte

community diversity index (CDI) for the created oxbow was similar, 1.61 and 1.65

respectively, during year 1 (the pulsing year) and year 3 (the year in which pulsing

returned) and lowest, 1.46, during the steady-flow year 2 (Table 4.2). The change in CDI

had strong correlations with the percent cover of the wetland (r2 = 0.97; Fig. 4.4a) and the

R-B flashiness index of the wetland (r2 = 0.98; Fig. 4.4b).

4.4.3 Macrophyte productivity

Total macrophyte production of the wetland decreased by 15.5 % from 7,560 kg yr-1 to 6, 544 kg yr-1 after the pulsing was removed (Fig. 4.5). This decrease was a result

of the areal macrophyte coverage within the wetland dropping from 32% to 25%. When

pulsing hydroperiod conditions were restored in the third year, the areal coverage

returned, 32%, and total net primary productivity increased 35% to 9,916 kg yr-1. There

107 was a positive correlation (r2 = 0.75) between the R-B flashiness index and the total macrophyte production, suggesting that flood pulsing may be a key forcing function for productivity (Fig. 4.4c).

The largest percent component of the total biomass production each year was

Typha spp. (Table 4.3). The total production of Typha decreased slightly (3%) when pulsing was removed but increased substantially (46%) when it was restored (Fig. 4.6).

The second most productive community was the mixed macrophyte community. The mixed macrophyte community, comprised primarily of Eleocharis sp., Scirpus americanus, Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis, decreased significantly in total production (15%) compared to the previous year when pulsing was removed but significantly increased (13%) compared to the previous year when pulsing was restored (Figs. 4.3, 4.6). The decrease in the percentage of the total macrophyte biomass comprised by the mixed community from

20.2% to 14.3% when pulsing was restored (Table 4.3) is an artifact of the large increase in Typha production and not indicative of a loss in mixed community production.

Phragmites australis, an invasive but native plant in this region, increased in total production each year of this study, doubling its production each year (Fig. 4.6). The total production of Sparganium eurycarpum decreased dramatically (91%) when pulsing was removed and recovered somewhat (73% of original total) when pulsing restored

(Fig. 4.6). Total production of Pontederia cordata increased by in each year of the study,

38% when pulsing was removed and 58% when pulsing was restored (Fig 4.6).

108 4.5 Discussion

4.5.1 The Effects of Pulsing

The springtime hydrologic pulses bring in nutrients (N-P), propagules, and fresh

sediments but the high water can also cause oxygen deprivation in the substrate and

impede macrophyte germination and growth. Other studies on pulsing suggest that some intermediate amount of pulsing maximizes ecosystem function (e.g. Connell, 1978;

Mitsch and Ewel, 1979; Odum et al., 1995; Townsend, 1996; Chapter 3). This theory suggests that wetland ecosystems with either a low or high degree of pulsing may provide fewer ecosystem services than wetland ecosystems with an intermediate degree of pulsing (Fig. 4.7).

The difference between years in this study was not only the amount of water flux into the wetlands (it ranged from 19 – 27 m yr-1 in this study), but also the timing of the

water delivery. This is apparent in the differences in the average monthly Richardson-

Baker flashiness indices between the flood pulse and steady-flow years. The index

during the flood pulse years, 0.96 ± 0.06 and 0.84 ± 0.06, was higher than the index

during the steady-flow year, 0.21 ± 0.02, suggesting that the wetland hydroperiod was

variable during the flood pulse conditions. This difference in the timing of water delivery

impacted the loading rate of the various nutrients into the wetland when the wetland held

water (Chapter 3), both of which likely had an impact on macrophyte growth.

The overall changes in net primary productivity in the wetland were the result of a

combination of changes in areal plant coverand unit area productivity. During steady-

flow hydroperiod conditions (year 2; 2005), the mean net aboveground primary

productivity per square meter of vegetation (NAPP) was 909 g m-2 yr-1. This was not

109 significantly higher (p = 0.10) than the 814 g m-2 yr-1 measured in the preceding pulsing

year (year 1; 2004), but it was significantly less (p = 0.05) than the 1111 g m-2 yr-1

measured in the following pulsing hydroperiod year (year 3; 2006; Table 4.4).

The productivity per unit area of Typha increased each year (Table 4.4) and was consistent in magnitude with other studies (Dubbe et al., 1988; Mason and Bryant, 1975;

Anderson and Mitsch, 2005). The percent composition of the total biomass also increased each year for Typha (Table 4.3). The reason the total production of Typha over

the entire wetland basin did not change significantly following the removal of river

pulsing (Fig. 4.6) was because of a reduction in its percent cover (Table 4.2).

The mixed macrophyte community also decreased in areal coverage when flood

pulsing was removed and increased when flood pulsing was restored (Table 4.2). The

opposite was observed for the productivity per unit area of the mixed macrophyte

community, with the steady flow year being the most productive per unit area. The

mixed community was the only community to have it’s greatest per unit area productivity

occur during the steady-flow year. It was, however, like each of the other observed

communities in that the greatest total production across the entire wetland basin occurring

during one of the pulsing hydsroperiod years. Part of this was driven by Cyperus

strigosis, being the only macrophyte species with a significant increase in presence (by

abundance, mass, and cover) during steady-flow compared to pulsing conditions. C.

strigosis grew in the outflow swale of the wetland outside of the inundated soil zone

during the steady flow year, but was insignificant during pulsing conditions in terms of

NAPP, total biomass, and percent cover.

110 P. australis, an invasive but native plant in this region, increased in per unit area

productivity and percent cover each year of the study (Tables 4.2, 4.3). Most of the P.

australis patch expansion was along the shore of the wetland above the inundated root

zone, with only a very small expansion within the inundated zone (Fig. 4.3). It appears

unlikely that the variation in hydroperiod had an effect on P. australis in this study.

S. eurycarpum had a large and significant decrease in areal coverage during the steady-flow year (Fig. 4.3). While it did recover somewhat following the restoration of pulsing conditions and even though the unit area net productivity increased significantly following the restoration of pulsing, the total production remained less than one-third of what it was prior to the removal of pulsing conditions (Table 4.4; Fig. 4.6).

The per unit area net productivity of P. cordata decreased with the onset of

flooding, but rebounded when flooding was restored. There was a steady increase in the

coverage of P.cordata throughout the course of the study.

4.5.2 Causes for variation in macrophyte productivity

The differences in total macrphyte productivity between years is likely a result of

the differences in hydrology. Productivity was high during a normal river-pulsing year,

decreased when the pulses were removed, and then rebounded to the highest productivity

of the study period when flood pulsing resumed. The pulsing hydroperiod creates

temporal variation in potential macrophyte habitat. The variation in pulsing frequency

from year to year (inter-annual comparison) of this study may have provided even more

“hydrologic diversity” over the three years that led to the highest productivity in the third

year.

111 As the water level rises and falls with the inflowing pulses, not only does the depth of the wetland change, but the areal extent of the wetland also changes as the shallowly sloped edges of the wetland are alternately covered by water and exposed to the atmosphere. Under a pulsing hydroperiod, this fluctuating littoral zone supports a variety of emergent macrophytes. During the steady-flow year, a lack of draw-down conditions and more deepwater conditions may have inhibited germination and reduced growth along this edge, leading to less areal coverage and reduced.

Soil exposure, which occurs during pulse draw-downs, has been shown to be positively correlated with species richness and species composition (Atkinson et al.,

2005). In the oxbow wetland in our study, we saw a similar correlation between R-B flashiness (which can lead to pulse draw-downs) and both community diversity and percent cover. In the steady flow year, water remained too deep in many areas for emergent macrophytes to germinate. The areal coverage of water in the wetland was also static. Steady-flow meant that there was no exposed mudflat during the dry season, which is an area of the wetland that colonized with macrophytes between floods during the pulsing hydroperiod years.

There was also a shift in the dominance of Typha spp. when flood pulsing was removed and when it was returned (Table 4.3). After a decrease in the percentage of the biomass comprised of Typha from 2003 to 2004 reported in another study (Fink and

Mitsch, in press), the percent Typha cover increased in 2005 during steady-flow conditions. This is possibly due to the static water levels along the edge of the open water portion of the wetland remaining too deep to provide suitable habitat for competing macrophyte germination (Keddy and Reznicek; 1986, Squires and van der Valk, 1992).

112 It is also possible that this could be a successional trend as It is also possible that this

could be a successional trend as Typha dominance may be increasing through clonal

vegetative spread (Fennesy and Cronk, 2001). This is the section of the wetland that had

the greatest amount of non-Typha macrophytes during the pulsing hydroperiod

conditions. The increase from 2004 to 2005 may also have been a result of the shallower

than typical water depths (1.85 ± 0.43 ft in 2004, 1.41 ± 0.08 ft in 2005; p < 0.001) in the spring and early summer allowing for greater Typha spread.

A decrease in total emergent macrophyte production (kg yr-1) during non-pulsing

hydroperiod conditions in the created oxbow is the opposite of what was observed by

Mitsch et al. (2005b) who observed 46% and 81% decreases in productivity with the

onset of artificial flood pulsing in two pumped wetlands at the same wetland research

park. Mitsch et al. (2005b) also reported a decrease in macrophyte diversity in a pumped

wetland experiment during pulsing conditions, again, the opposite of what was observed

in the oxbow wetland in this study. These differences between the results of our study

and the Mitsch at al. (2005b) study may have to do with the experimental design. In their

study, the river was flooded during the first week of every month during the wet season,

no matter what the stage of the adjacent river. In our study of the created oxbow, the

pulsing was caused by rises in river stage. Flow rates in rivers in the Midwest United

States are often correlated with nutrient loadings, especially during the wet season during

the spring and early summer following snow melt and farm field fertilization. Therefore

at the same time as the plants in the oxbow were receiving additional water stress, they

were also receiving nutritional subsidies (Fink and Mitsch, in press) whereas the plants in

the Mitsch et al. (2005b) study did not necessarily receive this subsidy.

113 In a mesocosm experiment, Anderson and Mitsch (2005) found that there was no significant difference in peak dry-weight biomass between pulsed and steady-flow conditions for Typha spp. or for Schoenoplectus tabernaemonti. The study did show that

Typha was more resilient to prolonged hydroperiods than other emergent macorphytes.

This finding does corroborate with our observation in the created oxbow that the per-unit area NAPP of Typha was less negatively affected by the removal of pulses than were most of the other emergents, especially those in the mixed macrphyte community (e.g

Schoenoplectus tabernaemontani, Sparganium eurcarpum, Juncus effusus, Scirpus americanus; Table 4.4). Other studies have reported that the rhizomes of S. tabernaemontani do not tolerate prolonged high water levels and the concordant anoxic conditions (van der Valk and Davis, 1978). This may partially explain why the lowest observed S. tabernaemontani productivity in this study was in the steady-flow year in which the water depth was consistent and there was very little fluctuating edge on which the S. tabernaemontani could colonize and thrive. The 1-m2 mesocosms in the Anderson and Mitsch (2005) study received 10.7 m-2 yr-1 of water inflow during the growing season, or about half the loading rate in our study of the created oxbow. This difference in hydraulic loading rate may explain the differences seen between the small-scale and the whole ecosystem scale study. Another possibility is that in the mesocosm study there was likely less drying of the soils due to reduced seepage and lowered evaporation which are common potential drawbacks to mesocosm studies (Ahn and Mitsch, 2001). This would have led to greater soil saturation in the root zone of the macrophytes than in the full ecosystem study, and may have inhibited effects associated with pulsing (e.g. mineralization). Further research on this topic may be necessary.

114

4.5.3 Nutrients and wetland vegetation patterns

The extent of the emergent marsh section of the created oxbow corresponds to the area of the marsh that had enriched nitrate concentrations as described by Fink and

Mitsch (in press) and in Chapter 3. There is a correlation (r2 = 0.89) between the spatial pattern of Typha productivity in the emergent marsh section of the oxbow wetland and the decrease in nitrate concentration reported in Chapter 3 (Figs. 3.4, 4.8). This relationship suggests that the pulses of NOx-N from the river have a fertilizing effect on the production of Typha specifically and on the macrophytes in general. Several studies have reported an increase in organic-N concentration in the vegetated portions of riparian wetlands that are nitrogen enriched like this oxbow wetland, followed by a reduction in organic-N after the NOx-N concentration is reduced (Lane et al., 1999; Chapter 3).

4.5 4 Management implications

Total cover requirements in compensatory mitigation projects are often set at 75% or 85% plant coverage for the first two years (Campbell et al., 2002). As such, the created oxbow in this study could have easily been viewed as a failed project (25-33% macrophyte coverage) despite its high level of ecosystem function (Fink and Mitsch, in press; Chapter 3; this study). This diversion wetland had a greater areal extent of macrophyte vegetation during the pulsing hydroperiod years than the steady-flow hydroperiod year. A pulsing hydroperiod should enable mitigation wetlands to reach target macrophyte coverage goals more easily. Nevertheless, this wetland is not designed to have 70 to 80% plant cover; the natural hydrologic pattern of high water in the spring

115 and low water in the fall in Midwestern USA riverine wetlands like this one will never have high plant coverage except for years with exceptional drought periods in the spring.

This wetland has not experienced those conditions since it was created in 1996.

4.6 Acknowledgements

Support provided by U.S. Department of Agriculture NRI CSREES Award 2003-

35102-13518 and a Payne grant from the Ohio Agricultural Research and Development

Center of The Ohio State University. We also thank Colleen Fink, Cheseaquah Blevins,

Andrew Tweel, Brittany Cleveland, Paul Jones, Ryan Younge, and Dr. Li Zhang for their help in biomass sampling.

116 4.7 Literature Cited

Ahn, C. and W.J. Mitsch. 2001. Scaling considersations of mesocosm wetlands in simulating large freshwater marshes. Ecological Engineering 18:327-342.

Anderson C.J., and W.J. Mitsch. 2005. Effect of pulsing on macrophyte productivity and nutrient uptake: a wetland mesocosm experiment. American Midland Natualist 154:305-319.

Atkinson, R.B., J.E. Perry, and J. Cairns Jr. 2005. Vegetation communities of 20-year- old created depressional wetlands. Wetlands Ecology and Management 13:469–478.

Baker, David B., R. Peter Richards, Timothy T. Loftus, and Jack W. Kramer. 2004. A New Flashiness Index: Characteristics and Applications to Midwestern Rivers and Streams. Journal of the American Water Resources Association 40:503-522.

Brown, S.L. 1981. A comparison of the structure, primary productivity, and transpiration of cypress ecosystems in Florida. Ecological Monographs 51:405-415.

Campbell, D.A., C.A. Cole, and R.P. Brooks. 2002. A comparison of created and natural wetlands in Pennsylvania, USA. Wetland Ecology and Management 10:41-49.

Casanova, M.T and M.A. Brock. 2000. How do depth, duration, and frequency of flooding influence the establishment of wetland plant communities. Plant Ecology 147:237-250.

Connell, J.H. 1978. Diversity in tropical rainforest and coral reefs. Science. 199:1302-1310.

Cronk J.K. and W.J. Mitsch. 1994. Periphyton productivity on artificial and natural surfaces in four constructed freshwater marshes under different hydrologic regimes. Aquatic Botany 48:325-342.

Day, J. W., T. J. Butler, W. G. Conner. 1977. Productivity and nutrient export studies in a cypress swamp and lake system in Louisiana. In M. Wiley, ed. Estuarine Processes, Vol. II. Academic Press, New York, pp. 255-269.

Day J.W., D. Pont, P.F. Hensel, C. Ibanez. 1995. Impacts of sea-level rise on deltas in the Gulf of Mexico and the Mediterranean: the importance of pulsing to sustainability. Estuaries 18: 636–647.

Day, J.W., L.D. Britsch, S. Hawes, G. Shaffer, D.J. Reed, and D. Cahoon. 2000. Pattern and process of land loss in the Mississippi Delta: a spatial and temporal analysis of wetland habitat change. Estuaries 23:425-438.

117 Dubbe, D.R., E. G. Graver, and D.C. Pratt. 1998. Production of cattail (Typha spp.) biomass in Minnesota, USA. Biomass 23:93-185.

Fennessy, M.S., and J. K. Cronk. 2001. Wetland Plants: Biology and Ecology. CRC Press.Boca Raton FL.

Fink, D.F. and W.J. Mitsch. in press. Hydrology, biogeochemistry, and plant community development in a created river diversion oxbow wetland. Ecological Engineering.

Galat D.L., L.H. Frederickson, D.D. Humburg, K.J. Bataille, J.R. Bodie, J. Dohrenwend. 1998. Flooding to restore connectivity of regulated, large-river wetlands. (Lower Missouri River). BioScience 48: 721–733.

Gathumbi, S.M., P.J. Bohlen, D.A. Graetz. 2005. Nutrient enrichment of wetland vegetation and sediments in subtropical pastures. Soil Science Society of America Journal 69:539-548.

Giovanni, S.GT. and D.M.L. Da Motta Marques. 1998. Establishment of three emergent macrophytes under different water regimes. Water Science and Technology 40:233-240.

Gusewell, S., U. Bollens, P. Ryser, F. Klotzli. 2003. Contrasting effects of nitrogen, phosphorus and water regime on first- and second-year growth of 16 wetland plant species. Functional Ecology 17:754-765.

Hein, T., G. Heiler, D. Pennetzdorfer, P. Riedler, M. Schageri, and F. Schiemer. 1999. The Danube Restoration Project: functional aspects and plantonic productivity in the floodplain system. Regulated Rivers 15:259-279.

Henry, C.E., C. Amoros, N. Roset. 2002. Restoration ecology of riverine wetlands: a 5- year post-operation survey on the Rhone River France. Ecological Engineering 18:543-554.

Keddy, P. A., and A. A. Reznicek. 1986. Great Lakes vegetation dynamics: the role of fluctuating water levels and buried seeds. Journal of Great Lakes Research 12, 25–36.

Kellogg, C.H., S.D Bridgham, S.A. Leight. 2003. Effects of water level , shade, and time on germination and growth of freshwater marsh plants along a simulated successional gradient. Journal of Ecology 91:274-282.

Lane, R.L., J.W. Day, D. Justica, E. Reyes, B. Marx, J.N. Day, E. Hyfield. 2004. Changes in stoichiometric Si, N and P ratios of Mississippi River water diverted through coastal wetlands to the Gulf of Mexico. Estuarine, Coastal and Shelf Science 60:1-10.

118 Mason, C.F. and R.J. Bryant. 1975. Production, nutrient content, and decomposition of Phragmites communis, Trin. and Typha angustifolia L. Journal of Ecology 63:71-95.

McDougal, R.L., L.G. Goldsborough, B.J. Hann. 1997. Responses of a prairie wetland to press and pulse additionsl of inorganic nitrogen and phosphorus: Production by planktonic and benthic algae. Archiv für Hydrobiologie 140: 145-167.

Megonigal, J. P., W. H. Conner, S. Kroeger, R. R. Sharitz. 1997. Aboveground production in southeastern floodplain forests: A test of the subsidy-stress hypothesis. Ecology 78:370-384.

Mitsch, W.J. and K.C. Ewel. 1979. Comparative biomass and growth of cypress in Florida wetlands. American Midland Naturalist 101:417-426.

Mitsch, W.J., C.L. Dorge, J.W. Wiemhoff. 1979. Ecosystem dynamics and a phosphorus budget of an alluvial cypress swamp in southern Illinois. Ecology 60:1116-1124.

Mitsch, W.J. and W.G. Rust. 1984. Tree growth responses to flooding in a bottomland forest in northeastern Illinois. Forest Science 30:499-510.

Mitsch, W.J., J.R. Taylor, K.B Benson. 1991. Estimating primary productivity of forested wetland communities in different hydrologic landscapes. Landscape Ecology 5:75-92.

Mitsch, W.J., N. Wang, L. Zhang, R. Deal, X, Wu, A. Zuwerink. 2005a. Using ecological indicators in a whole-ecosystem wetland experiment. pp.211-235. In: Jorgenson, S.E., F.L. Xu, R. Costanza (Eds.), handbook of Ecological Indicators for Assessment of Ecosystem Health, CRC Press, Boca Raton, FL.

Mitsch, W.J., L. Zhang, C.J. Anderson, A.E. Altor, M.E. Hernandez. 2005b. Creating riverine wetlands: Ecological succession, nutrient retention, and pulsing effects. Ecological Engineering. 25:521-527.

Mitsch, W.J. and J.W. Day. 2006. Restoration of wetlands in the Mississippi-Ohio- Missouri (MOM) River Basin: Experience and needed research. Ecological Engineering 26:55-69.

Newman, S., J. Schuette, J.B. Grace, K. Rutchey, T. Fontaine, K.R. Reddy, M. Pietrucha. 1998. Factors influencing cattail abundance in the northern Everglades. Aquatic Botony 60:265-280.

Nixon, S.W. 1988. Physical energy inputs and the comparative ecology of lake and marine ecosystems. Limnology and Oceanography 33:1005–1025.

Odum, E.P. 1990. Field experimental tests of ecosystem-level hypotheses. Trends in Ecological Evolution 5:204–205.

119 Odum W.E., E.P. Odum, H.T. Odum. 1995. Nature’s pulsing Louisiana. American Association of Petroleum Geologists Bulparadigm. Estuaries 18: 547–555.

Olila, O.G, K.R. Reddy, D.L. Stites. 1997. Influence of draining on soil phosphorus forms and distribution in a constructed wetland. Ecological Engineering 14, 107-126.

Reed, P.B. Jr. 1998. National list of plant species that occur in wetlands: Northeast (Region I). U.S. FWS, Washington, DC, Biological Report 88 (26.1).

Reyes, E, J.F. Martin, J.W. Day, G.P. Kemp, H. Mashriqui. 2004. River forcing at work: ecological modeling of prograding and regressive deltas. Wetlands Ecology and Management 12:103-114.

Spink, A., R.E. Sparks, M. van Oorschot, and T.W. Verhoenven. 1998. Nutrient dynamics of large river floodplains. Regulated Rivers: Research and Management 14:203-216.

Squires, L. and A.G. van der Valk. 1992. Water-depth tolerances of the dominant emergent macrophytes of the Delta Marsh, Manitoba. Canadian Journal of Botany 70:1860-1867.

Tanner, C.C., J. D’Eugenio, G.B. McBride, J.P.S. Sukias, K. Thompson. 1999. Effect of water level fluctuation on nitrogen removal from constructed wetland mesocosms. Ecological Engineering 12:67-92.

Townsend, C.R. (1996) Concepts in river ecology: pattern and process in the catchment hierarchy. Archiv fur Hydrobiologie Supplement 113, Large Rivers 10: 3-21

Tuttle, C.L. and W.J. Mitsch. in review. Aquatic metabolism as an indicator of the ecological effects of hydrologic pulsing in flow-through wetlands. Ecological Indicators

United States Bureau of Reclamation (1997). Water Measurement Manual 3ed. U.S. Department of the Interior, Bureau of Reclamation. Washington D.C. van der Valk, A.G. and C.B. Davis. 1978. The role of seed banks in the vegetation dynamics of prairie glacial marshes. Ecology 59:322-335.

Waters, I. and J.M Shay. 1992. Effect of water depth on population parameters of Typha glauca stands. Canadian Journal of Botany 70:349-351.

120

Figure 4.1 Site map of the Wilma H. Schiermeier Olentangy River Wetland Research

Park on The Ohio State University campus. The 2.8 ha created oxbow wetland is between the experimental wetlands and the bottomland hardwood forest. The locations of the seven biomass sampling transects are marked with white lines.

121 122

Figure 4.2 Hydrograph for the created oxbow wetland and the Olentangy River from

April 2004 to September 2006.

123

124

Figure 4.3 Macrophyte communities in the created oxbow wetland during pulsing (2004

and 2006) and steady flow (2005) conditions during peak biomass. The area in the 2004

map marked with dotted lines shows the spread of Xanthium strumarium following a prolonged drawdown that occurred after the biomass surveys were completed.

125

0 200 feet 0 60 meters

- Xanthium strumarium N 126

126

Figure 4.4 Comparisons of: (a) the community diversity index and percent areal coverage of emergent macrophytes (r2 = 0.97), (b) the the community diversity index and the

Richardson-Baker flashiness index (r2 = 0.98), and (c) the total macrophyte production of the wetland and the Richardson-Baker flashiness index (r2 = 0.75).

127 1.70 2006 - pulsing 1.65 2004 - pulsing 1.60 y = 0.0263x + 0.7849 R2 = 0.9547 1.55

1.50

Community Diversity Index . 2005 – steady-flow 1.45 25 27 29 31 33 Percent Cover

1.70 2006 - pulsing 1.65 2004 - pulsing 1.60 y = 0.2755x + 1.4016 R2 = 0.9703 1.55

1.50

Community Diversity Index . 2005 – steady-flow 1.45 0.0 0.2 0.4 0.6 0.8 1.0 R-B Index

12000 y = 3627.8x + 5745.3 2006 - pulsing R2 = 0.564 )

-1 9000 2004 - pulsing 6000 2005 – steady-flow

Total macrophyte 3000 production (kg yr production

0 0.0 0.2 0.4 0.6 0.8 1.0 R-B Index

128

Figure 4.5 Total net aboveground primary productivity in the created oxbow wetland during pulsing (2004 and 2006) and steady flow (2005) during peak biomass. Labels a and b indicate statistical differences in the mean biomass between years (p = 0.36 between 2004 and 2005; p = 0.05 between 2004 and 2006; p = 0.02 between 2005 and

2006). Error bars indicate standard error.

129

b

a

a 130

130

Figure 4.6 Total production (kg yr-1) of the 5 dominant macrophyte communities in the created oxbow wetland in 2004 (pulsing hydrology), 2005 (steady flow hydrology), and 2006 (restored pulsing hydrology). The “mixed macrophyte” community is comprised predominately of:

Eleocharis sp., Scirpus americanus, Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis.

131

132

132

Figure 4.7 Effects of pulsing on wetland ecosystem services. Arrows indicate the direction of increased service for each process. A bi-directional arrow indicates uncertainty as to the effect of pulsing with the relative size of the arrowhead suggesting the general impact.

133

Effect Pulsing Pulsing removed

Hydroperiod R-B flashiness HRT Vegetation Percent cover NAPP Diversity Total Production Animals Avian use Nutrients (N&P) Loading rate Retention rate Percent removal Suspended Solids Loading rate Retention rate Percent removal

134

Figure 4.8 Pattern of Typha net above ground primary productivity per unit area

(NAPP, g dry weight m-2 yr-1) compared to the nitrate + nitrite concentration in the surface water from the inflow, through the emergent marsh, open water, and outflow of the created oxbow wetland.

135

136

136

Pulsing Steady-flow Pulsing 2004-2005 2005-2006 2006* Wet Season Days of flow 165 148 25 Inflow m3 m-2 19 11 8 Outflow m3 m-2 17 11 7 HRT d 6 ± 1 14 ± 1 9 ± 1 Mean depth m 1.40 ± 0.42 1.39 ± 0.11 1.14 ± 0.43 Dry Season Days of flow 7 90 10 Inflow m3 m-2 8 9 5 Outflow m3 m-2 9 9 5 HRT d 4 ± 1 7 ± 1 6 ± 1 Mean depth m 1.11 ± 0.22 1.35 ± 0.08 1.11 ± 0.41 Annual Number of Pulses 8 0 6 Total Days of flow 172 338 35 Inflow m3 m-2 27 20 13 Outflow m3 m-2 26 20 12 HRT d 6 ± 1 12 ± 1 8 ± 0 Mean depth m 1.32 ± 0.40 1.38 ± 0.10 1.13 ± 0.42 R-B index 0.82 ± 0.06 0.21 ± 0.02 0.84 ± 0.06

Table 4.1 Inflow, outflow, mean hydraulic retention time (HRT; ave ± std error), and

water depth (ave ± std dev) of the mitigation oxbow wetland in April 2004 – Sept 2006.

Units of inflow and outflow are cubic meters of water flux per square meter of wetland

area (2.8 ha). HRT was only calculated for days in which there was inflow or outflow.

The Richardson-Baker (R-B index) flashiness index for the created oxbow wetland was calculated on a monthly basis for the pulsing and steady-flow hydrologic conditions. The

Wet Season is defined as November through June; the Dry Season is defined as July through October. *The second pulsing year (2006) only encompasses a partial year (170 days, April 2006 to September 2006).

137

2004 2005 2006 Typha spp. 14.6% 12.1% 16.0% Sparganium eurycarpum 2.6% 0.3% 0.5% Pontederia cordata 0.2% 0.4% 0.7% Phragmites australis 0.8% 1.2% 1.9% Mixed macrophytes 14.1% 11.7% 12.9% Total coverage 32.3% 25.7% 32.1%

CDI = Σ Ciln(Ci) 1.61 1.46 1.65

Table 4.2 Percent areal coverage and the Community Diversity Index (CDI) of the 5 dominant macrophyte communities in the created oxbow wetland in 2004 (pulsing hydrology), 2005 (steady flow hydrology), and 2006 (restored pulsing hydrology). The

“Mixed macrophyte” community is comprised predominately of: Scirpus americanus,

Eleocharis sp., Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis. NAPP is calculated for the area where macrophytes actually grew and not for the area of the entire oxbow wetland. CDI = Σ (Ciln(Ci)). Where Ci is the present cover of community “i” and N is the number of macrophyte communities (Mitsch et al., 2005a).

138

2004 2005 2006 Typha spp. 71.2% 74.8% 76.8% Sparganium eurycarpum 6.2% 0.8% 1.6% Pontederia cordata 0.3% 0.4% 0.7% Phragmites australis 1.6% 3.7% 6.6% Mixed macrophytes 20.7% 20.2% 14.3% Total biomass 100.0% 100.0% 100.0%

Table 4.3 Percent of the total biomass comprised by each of the 5 dominant macrophyte communities in the created oxbow wetland in 2004 (pulsing hydrology), 2005 (steady flow hydrology), and 2006 (restored pulsing hydrology). The “mixed macrophyte” community is comprised predominately of: Eleocharis sp., Scirpus americanus, Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis.

139

-2 -1 Net aboveground Primary Productivity (g m yr ) 2004 2005 2006 Typha spp. 1305 ± 150 (5)a 1580 ± 134 (5)b 1694 ± 115 (6)c d d e Sparganium eurycarpum 640 ± 32 (2) 646 ± 32 (2) 1085 ± 54 (2) f g f Pontederia cordata 368 ± 18 (2) 260 ± 13 (2) 342 ± 17 (2) h i j Phragmites australis 549 ± 27 (2) 797 ± 40 (2) 1214 ± 61 (2) k k k Mixed macrophyte 393 ± 55 (6) 442 ± 24 (4) 394 ± 82 (5) l l m Mean 814 ± 54 (17) 909 ± 58 (16) 1111 ± 62 (17) (average ± standard error (number of samples)

Table 4.4 Net aboveground primary productivity per unit area of vegetation (NOT total

production) for the 5 dominant macrophyte communities in the created oxbow wetland in

2004 (pulsing hydroperiod), 2005 (steady-flow hydroperiod), and 2006 (restored pulsing

hydroperiod). Superscripts denote significance between years for each macrophyte

community (p < 0.05). The “mixed macrophyte” community is comprised predominately

of: Eleocharis sp., Scirpus americanus, Juncus effusus, Leersia oryzoides, Sagittaria spp., Verbesnia alterniflora, and Cyperus strigosis.

140

CHAPTER 5

USING SHORT-TERM HYDROLOGIC DATA TO PREDICT LONG-TERM

SUCCESIONAL DEVELOPMENT IN CREATED RIPARAIN WETLANDS

5.1 Abstract

The objective of this study was to create a model that could use short-term hydrologic data to predict long-term successional development in a created riparian wetland ecosystem. The hydraulic submodel predicted wetland water depth based on precipitation, potential evapotranspiration, river stage, overland outflows, and groundwater exchange. Vegetation submodels calculated the growth of trees and emergent macrophytes based water depth, flood duration and timing, and land elevation.

The model predicted the state of the ecosystem as open water, mudflat, marsh, or forested wetland. Results of the vegetation submodel at specific land elevations were then applied spatially to predict long-term dynamics for a created riparian wetland located at the

Schiermeier Olentangy River Wetland Research Park in central OH, USA. Data from

2004 were used to calibrate the hydrologic and vegetation submodels. Simulation results were validated with field data from 2003, 2005, and 2006. When the wetland inflow was flashy, the model predicted water depth within 6 cm of actual data 75% of the time; when

141

it was steady in flow model results were within 3 cm of actual data 75% of the time.

Simulations of vegetation succession were evaluated at 8 (current wetland age) and 100 years. Simulations for the current wetland age correctly predicted the vegetation patterns observed in the ecosystem. Simulations of wetland succession including a sediment accretion subroutine predict hydrarch succession, with forested wetland eventually replacing emergent marsh and open water areas of the wetland. Model function could potentially be improved by adding a dispersal model for propagules, a shorter time-step for vegetation, and an advective flow component to the hydrologic model.

5.2 Introduction

Both autogenic (internal biotic) and allogenic (external physical) processes are important in the pathway of development and the final characteristics of mature wetland ecosystems. In the classical view of succession, wetlands are considered transient stages in the hydrarch development of a terrestrial forested climax community from an open water ecosystem (Mitsch and Gosselink, 2000). Primary succession of plant communities directed towards a climax; however, is not a typical occurrence in many wetlands because these ecological systems are inherently dependent on changes in hydroperiod. Temporal hydrological variability often causes reversals or setbacks in succession (Fig. 5.1). In these wetland types (e.g. tidal marshes, riparian oxbows, seasonal prairie potholes), hydrologic variability dictates that allogenic processes typically overwhelm autogenic processes (Wilcox, 2004).

In semi-permanent wetlands and seasonal wetlands (Cowardin et al., 1979; Mitsch and Gosselink, 2000), wet or dry periods induce changes in the relative amount of

142

emergent cover, mudflat, and open water (Poiani and Johnson, 1993). Timing and duration of wet and dry periods also influences coverage of herbaceous and woody vegetation in riparian wetlands (Bedinger, 1971; Jowarski et al., 1979; Karr, 1989; Keddy and Reznicek, 1986; Johnson, 1994; Toner and Keddy, 1997; Deberry and Perry, 2004).

Newly exposed hydric soils are colonized primarily by annual species in the first season of exposure and then dominated by perennial species in following years (Noon, 1995).

As wet conditions return, annuals are eliminated and perennial emergents are stimulated to spread vegetative (van der Valk and Davis, 1978). During this phase the wetland is dominated by emergent macrophytes. Open water areas increase as emergents in the deeper part of the basin are eliminated because of prolonged flooding and emergent cover is then reduced to a narrow landward band around the fringe of the wetland (Poiani and

Johnson, 1993). The structure and species composition of riparian wetland vegetation constitute fundamental elements of the diversity of riparian ecosystems (Nilsson, 1992;

Malanson, 1993). These assemblages are generally high in species numbers (Nilsson et al., 1988; Nilsson, 1992) and encompass structurally distinguishable vegetation types, from trees to herbaceous vegetation (Nilsson, 1984, 1992).

Rapid changes in community structure may be relatively easy to quantify, but long-term changes are more subtle and difficult to predict, requiring simulation modelling to examine the mechanisms and complex interactions affecting vegetation change

(Sturtevant, 1998). Using simulation models to examine ecosystem dynamics over long time scales may be of value as the revegetation of a restored or created wetland over a short period of time is no guarantee that the wetland will continue to function over time

(Kusler and Kentula, 1990; Atkinson et al., 2005).

143

Such process-oriented ecological models can be useful tools in ecosystem research and management. Ecosystem model objectives may range from those involving predictions on the scale of individual organisms or populations to models that focus more on quantifying interactions in ecosystems. Moreover, model objectives that are similar for different biomes or regions may involve different physical forcing functions or feedback mechanisms that significantly alter ecosystem behavior (Fitz et al., 1996).

Central to many generalized ecosystem models has been the assumption of homogeneity within the system, using lumped parameters. Some models have attempted to accommodate this short-coming by replicating unit models in a grid-cell array for explicit spatial simulations (e.g. Costanza and Maxwell, 1991; Poiani and Johnson, 1993;

Ellison and Bedford, 1995; Sturtevant, 1998; Martin et al., 2002). The authors in these papers were able to quantitatively simulate vegetation change in response to varying water levels by modelling life history traits as a mathematical function of water depth.

Hydrologic inputs are a critical component of many ecological systems (van der

Valk, 1981; Fitz et al., 1996). Chance events, such as flood pulsing or drought years, may affect the long-term composition of a wetland by allowing species establishment

(van der Valk, 1981; Middleton, 2000). Furthermore, tolerance to flooded conditions carries an energetic cost, so less flood-tolerant species are superior competitors in drier locations (Grace and Wetzel, 1981), allowing invasion of occupied space when water tables are low for extended periods of time. A potentially important effect that happens in the real world but that models often overlook is the influence that plant roots can have on evapotranspiration and on the distribution of soil water via the process of hydraulic lift. Deep-rooted trees take in water from deeper moist soil layers and exude that water

144

into the drier soil layers near the surface where it either evaporates directly out of the soil surface or is available for shallow rooted macrophytes to use and transpire (Chahaine,

1992; Feddes et al., 2001). Fluxes along the soil-plant-atmosphere continuum are regulated by above- and below-ground plant properties (Dawson et al., 1998; Jackson et al., 2000a,b). Which, in addition to the water use needs of the trees themselves, effectively increases the rate of potential evapotranspiration from the near surface soil itself.

5.2.1 Goals and Objectives

The goal of this research is to integrate short-term and long-term models to investigate potential trajectories and endpoints for the ecological succession of a created riparian river diversion wetland. The output of this model represents a square meter of wetland and can be transferred to a GIS for a spatial representation of a riparian wetland.

Pursuant to this goal this study had the following objectives.

1. Develop a dynamic unit model to predict the hydroperiod of a created oxbow

wetland based on annual of river flows, precipitation, and evapotranspiration.

2. Develop a unit model to predict plant succession, the ecological state (open water,

mudflat, marsh, forested wetland), and the effect of altered water levels and

sedimentation on vegetation growth.

3. Integrate the hydrologic and vegetation models to make long-term prediction

about the succession in the entire wetland over a 100-year period.

145

5.2.2 Site Description

The 3-ha wetland modelled in this study is located on the floodplain of the

Olentangy River in Central Ohio at the Schiermeier Olentangy River Wetland Research

Park at The Ohio State University (Columbus, Ohio, USA; Fig. 2.1). Water enters the

oxbow through a Red Field TideflexTM check valve when the river elevation is higher than the wetland, and flows back to the Olentangy River though an outflow control weir by. The wetland has two significant vegetation zones. The northern half (closest to the inflow) is an emergent marsh, and the southern half (closest to the outflow) is an open water basin. The wetland was constructed in 1997 and planted with 6900 rootstocks representing 21 species (Mitsch et al., 1998; Fink and Mitsch, in press). In addition, sapling and 12, 1.5 cm trees were planted along the fringe of the wetland in 2003. These plantings provided a small seed bank and a jumpstart on primary wetland succession.

5.3 Modeling Methods

The general modelling approach is followed the procedure described by Jørgensen

and Bendoricchio (2001). The one exception from this method was a validation step was

not conducted on some parts of the model due to limited available data. The model was

constructed using STELLATM (Structural Thinking, Experimental Learning Laboratory

with Animation) language and software (Richmond et al., 1987). The program is a

graphically based simulation environment that alleviates the need for being an expert in

high level programming language to develop new models or to understand existing

models (Costanza, 1987). Furthermore, because a STELLA model can be easily divided

into functional submodels, it will be easier to add complexity in the future by

146

incorporating additional submodels into the existing models framework. The model is designed as a unit model describing ecosystem development at a particular land elevation in a theoretical square meter of wetland. Results from the unit model will then be extrapolated over the entire wetland basin by assuming that all land with the same elevation contour will have the same ecosystem response.

Three models were built and linked for this project: a hydroperiod model, an emergent macrophyte vegetation (marsh) model, and a woody vegetation (forested wetland) model. A set of non-linear ordinary differential equations was used in each submodel. Needed parameters were taken from published literature, measured in the field, estimated from field data, or determined through model calibration. The model provides an annual snapshot of the general ecosystem state (open water, mudflat, marsh, or forested wetland; Table 5.1). The interactions between hydroperiod, climate, and vegetation used in this modelling effort are indicated by the Odum diagram (Odum, 1983;

Brown, 2004) in Fig. 5.2. Water depth within the wetland basin is primarily controlled by the forcing functions of river inflow (controlled by river stage), precipitation, and surface outflows, with forested wetland plants playing a minor role in increasing the effects of evapotranspiration. The ecosystem state (open water, mudflat, marsh, forested wetland) is controlled by the presence of a viable seed bank and by seasonal water depth.

The ecosystem selected for by various water depths is indicated by the vertical order in which they are positioned in Fig. 5.2. Open water habitat is created when the wetland’s water level is high and forested wetland habitat being created when the wetland’s water level is low.

147

5.3.1 Calibration

The model was calibrated by adjusting selected parameters in the model to obtain a best fit between the model calculations and field data. A stepwise calibration procedure

(Mitsch and Reeder, 1991) was used by first calibrating the hydroperiod submodel, then the woody vegetation submodel, and finally the emergent macrophyte submodel.

Hydroperiod and vegetation models were calibrated using river stage, wetland inflow, wetland depth, vegetation coverage, and macrophyte net above ground primary productivity data collected between April 2004 and March 2005. As each unknown parameter was calibrated, values previously calibrated parameters were not varied. With this technique, several unknown coefficients in the model were given values (Tables 5.2;

5.3; 5.5).

5.3.2 Sensitivity analysis

Sensitivity analysis was performed to aid in model calibration and to assess how responsive different parts of the model are to simulated environmental variations.

Parameters, forcing functions, initial values, and model equations were systematically changed to correspond to the selected state variable. The sensitivity S(p) of a parameter, is defined by Jørgensen and Bendoricchio (2001) as:

S(p) = (Δx/x)/(Δp/p) (4.1)

148

5.3.3 Integration and modelling techniques

The hydrologic submodel (Fig. 5.3; Section 5.4.1) was integrated using fourth order Runge-Kutta methods with a time step of 1 day. The output from this model provided seasonal water levels that could be subsequently imported and used as the hydrologic inputs for the vegetation models (Poiani and Johnson, 1993). For most simulations, a single year of Olentangy River stages (2004) was extrapolated over a 100- year time period.

Vegetation (Section 5.4.2) simulations used Euler’s integration technique with a time step of 3 months and simulation period of 100 years. Simulation results were used to predict macrophyte and tree growth, and general areal coverage at the end of 8 and 100 years. No calibration or validation of the woody vegetation models was attempted because of the lack of data for the site. The woody vegetation model has, however, been used by Niswander (1995) and validated by Gamble (2006) who found that this model on average predicted tree height within 70 cm, and tree diameter within 2.7 cm for eight species studied in a wetland in central Ohio (Gamble, 2006).

Most studies of flooding effect on tree growth have compared annual growth measurements with mean water levels, number of floods, or average river discharge per year (Keeland and Sharitz, 1997). This study tries improve model accuracy by modelling changes in plant growth on a quarterly (seasonal) basis. Trees that are adapted to wetland habitats have the ability to withstand transient flooded conditions. And while trees have been shown to have physiological and growth responses to short-term flooding (Hook and Brown 1973; Keely, 1979; Kozlowski, 1984; Keeland and Sharitz, 1997) a quarterly time step provides an accurate representation of tree growth with water depth as a forcing

149

function (Botkin, 1993). A quarterly time step should be long enough adequately models

tree growth and the still be short enough to model the effects of short-term hydrologic

pulses flowing through the wetland on emergent macrophytes.

5.4 Model Structure

5.4.1 Hydrologic submodel

The water volume in the wetland was modeled using the following equation:

dV/dt = Qin(t) + Pt(t)Aw + GWin– k2Qout – ET – GWout (4.2)

The hydrologic submodel (Fig. 5.3) has only one state variable, water volume (V), which

balances direct precipitation (Pt), pumped and flooded river inflows (Qin),

evapotranspiration (ET), groundwater exchange (GWin, GWout), and outflow (Qout) from the wetlands. All flows were measured or estimated based on field data collected from

April 2004 to April 2006. The discharge from the wetland is controlled by the elevation of a V-notch weir (Lw) placed at the outflow. Actual outflows were measured at different

water surface elevations over the outflow weir. The actual and modeled outflows (Qout)

were plotted versus depth over the weir (Dow) to assist in calibration of the model.

Equations that calculated water level (MSLW) at a given water volume (V) in the wetland

were determined for the wetland basin based on bathymetric maps (Wang et al., 1998).

Average daily temperatures for Columbus, Ohio were used to calculate potential

150

evapotranspiration according to Thownwaite’s equation (Chow, 1964). Daily precipitation and temperature data were collected on site with missed data points being supplied by the NOAA/OARDC weather station (Table 5.2).

5.4.2Vegetation submodels

Vegetation within the simulated wetland was categorized into two major growth forms, forested (trees) and marsh (emergent herbaceous macrophytes). The emergent macrophyte growth form represents a combination of macrophyte functional groups that range from species that are relatively tall and highly flood resistant to species that are more commonly found on the only semi-inundated edge. The tree growth form represents taller woody plants that have a lower flood tolerance, but that can have a significant impact on the ecosystem through hydraulic lift and shading.

The model is constructed in a similar way to Ellison and Bedford (1995) and

Sturtevant (1998) in that each vegetation type germinate, grows, and experiences mortality, all as a direct function of the hydrologic state. Temporally, each simulation

“year” is divided into a time step of four “seasons” (spring, summer, autumn, winter), with germination from a seed bank assumed to only occur during the spring. Apart from submergent vegetation, which is not in this model, most wetland species require an exposed seedbed to germinate (van der Valk, 1981; Welling et al., 1988), though certain emergents may potentially grow in shallow water (Welling et al., 1988; Baldwin et al.,

1996; Mitsch and Gosselink, 2000). Seed germination in the model is related to the hydrologic state by a community specific parameter that defines the depth at which plants

151

can germinate. It was assumed that each square meter of the wetland contains an unlimited number of propagules available for germination (Poiani and Johnson, 1993) and that these propagules are evenly distributed throughout the wetland.

5.4.2.1 Sediment accretion sub-routine

In some of the simulations (e.g section 5.4.1.4; Simulation 4 –sediment accretion), a sediment accretion sub-routine was added to the vegetation growth models. The sediment accretion model acted as a switch that checked during each time step whether or not there was flooding.

IF(WD) > 0) THEN (0.25*Srate) ELSE (0) (4.3)

Wherein if the water depth (WD) was greater than zero, than a uniform deposition of sediments occurred according to an annual sediment deposition rate (Srate). If water depth was not greater than zero, then no deposition occurred.

4.4.2.2 Tree growth submodel

The tree growth submodel was developed to predict the germination, growth, and spatial extent of woody vegetation in the created oxbow wetland (Fig. 5.4). The model assumes that only a single tree can grow per square meter of wetland and is based on

Botkin’s (1993) optimum tree growth equation:

dTD/dt = GTTD[1-(TDTH/DmaxHmax)]/[274+3b2TD-4b3TD] (4.4)

152

TD is the tree diameter and Dmax is the maximum possible tree diameter. TH and Hmax are the tree height (eq. 4.4) and the maximum tree height possible respectively. GT, b2, and

b3 are species-specific growth parameters (Table 5.3). DT is the depth to the water table.

The height for the selected tree species was calculated using the following

equation (Botkin, 1993):

2 TH = f(TD) = 137 + b2TD – b3TD (4.5)

The growth parameters (b2 and b3) were taken from Botkin (1993), Pearstine et al.

(1985), or calculated using species information from Petrides (1988) or Sargent (1933).

Parameter values for selected species whose growth could be modeled in the created

oxbow wetland are given in Table 5.4. The tree species used for this model was Populus

deltoides (eastern cottonwood). Mortality in this model can occur if the seasonal growth

is less than a given minimum (Niswander and Mitsch, 1995). By dividing the growth

model into four seasons, we were able to have flooding early in the growing season

inhibit colonization by woody vegetation (Gill, 1970). Growing degree days (DegD) were also expected to have an effect on tree growth (Niswander and Mitsch, 1995) and were calculated according to Botkin (1993). Water table depth was also modeled to have an affect on tree growth and a water factor was calculated using Phipps (1979) equation

(Table 5.3):

2 WF = 1 – 0.055(WD - TOD) (4.6)

153

Where TOD is the species –specific optimum water table depth. This equation was only used if the seasonal water level was below the soil surface. If the seasonal water depth was above the soil surface, then tree growth was assumed to be zero for that season. The total production of woody biomass (kg/tree; BM) was then estimated using the following equation from Jenkins et al. (2004).

β + (β x ln(T )) BM = e 0 1 D (4.7)

Where β0 and β1 are species-specific parameters from Jenkins et al. (2003) and TD is the diameter of the tree at breast height in cm (Table 5.3).

5.4.2.3 Impact of trees on hydroperiod

In this modelling effort, the increase in root uptake due to the hydraulic lift in forested wetlands is applied only to the vegetation growth models. Root water uptake was modeled as (Feddes et al, 2001)

-C x Dw U = e 1 (4.8)

Where Ci the coefficient of hydraulic lift and Dw is depth to the water table (Table 5.3).

154

5.4.2.4 Emergent macrophyte growth submodel

The emergent macrophyte growth model (Fig. 5.5) assumes that within each square meter of wetland there is a maximum potential biomass that can be produced. The model predicts what percentage of that biomass is produced as a function of the water depth within that square meter and is modeled as Ellison and Bedford (1995).

dMacB/dt = MacB(t) + (Herb_NAPP - Herb_Death) (4.9)

Where MacB is the amount of macrophyte biomass, Herb_NAPP is the increase in macrophyte biomass and Herb-Death is the loss of macrophyte biomass (Table 5.5).

Berb_NAPP and Herb_Death are controlled by species-specific growth and death parameters. Each plant has an optimum depth at which it germinates and subsequently grows. Below this optimum depth, growth declines linearly with a rising water table, reaching zero growth at a maximum depth tolerance for growth (Ellison and Bedford,

1995; Sturtevant, 1998). Herb_NAPP is equal to ta germination function (see below) multiplied by the proportion of the maximum macrophytes production that grows in a square meter plot decreases linearly with increasing water depth (Kozlowski et al., 1991;

Squires and van der Valk, 1992):

PropG = C2(1 – 0.02*WD) (4.10)

Growth is also controlled by a function relating water depth to mortality. This function assumes that once established, a plant is able to survive even as conditions deteriorate,

155

but eventually water stress becomes so unfavorable that a high proportion of the plants

are no longer able to survive. Growth is also controlled by a mortality function that

increases exponentially with increasing water depth with 100 percent mortality below a

maximum.

0.017*WD PropD= C30.013(e ) (4.11)

The constants C2 and C3 are species-specific constants relating water depth to growth and death rates (Tables 5.5 and 5.6).

Seed germination is related to water depth by a species-specific germination

parameter that determines whether or not a species can germinate in a cell with a given

water level. Seeds of species that can germinate only during periods of draw down will

not germinate if its water depth is > 0. Similarly, species that can germinate only during

flooding conditions will not germinate if water depth = 0. Species that can germinate

regardless of the presence or absence of water will germinate as long as the cell’s water

depth is < 20 cm (Shay and Shay, 1986; Ellison and Bedford, 1995, Sturtevant, 1998).

Macrophytes are modeled as a combination community of multiple vegetative

functional groups (Boutin and Keddy, 1993). Deepwater growth characteristics are used

for Typha and edge growth factors are for littoral species akin to Juncus or Carex species

(Shay and Shay, 1986). This allows macrophytes to grow in the inundated and non- inundated elevation zones of the wetland without needing to have a separate submodel for deepwater and littoral macrophytes (for an example of this approach see Sturtevant,

1998).

156

5.4.3 Ecosystem state determination

The model also predicts the state of the ecosystem during peak biomass. If there is no macrophyte biomass, no woody vegetation biomass, and the water level is at or below the surface of the sediments, then the model reports “mudflats.” If there is no vegetation but the water level elevation is higher than the elevation sediments, then the model reports “open water.” If there is macrophyte biomass, the model reports “marsh.”

If there is woody vegetation, then the model reports “forested.” If there are both macrophytes and woody vegetation growing in the same unit, the model assumes that the macrophytes are the dominant vegetation types and returns “marsh” until the trees reach a diameter greater than 1 cm at which point the model reports “forested.”

5.5 Results and Discussion

5.5.1 Simulations

In addition to a baseline wetland succession simulation, scenarios were selected that represent alternate potential inflow conditions, the effects of sediment accretion, and the impacts of ecosystem engineers. This provided a range of potential end points for the development of a mature wetland ecosystem. A suite of seven elevations representing a gradient from the lowest point in the created oxbow wetland up to the dry uplands that are only rarely flooded (less than once per decade) was chosen for these simulations

(Fig. 5.6)

.

157

5.5.1.1 Simulation 1 – baseline, 2004 hydroperiod

If the wetland is receives 100 years of 2004 Olentangy River flow, the wetland develops into a heterogeneous ecosystem with the predicted outcome completely dependant on model assumptions and/or wetland management decisions (Fig 5.7).

Assuming no net sedimentation over this time, the wetland is predicted to have an open water area, two small areas of exposed mudflat, a marsh in the inflow portion, and a ring of forested wetland fringing the entire basin (Fig. 5.7a). This long-term picture is similar to what actually occurred in 2004, except for the emergent vegetation seen in 2004 along the littoral contours being replaced by woody vegetation. While 2004 was wetter than most other recent years (Fink and Mitsch, in press) the pattern of river flow was typical for the Olentangy River and for temperate rivers in general (Vannote et al., 1980). As such 2004 hydrologic data were used as the baseline for each of the following simulations to investigate how changes in river flow, wetland management, or model assumptions would affect long-term wetland successional development.

5.5.1.2 Simulation 2 – dramatic flood pulsing

As rivers become more flashy (Baker et al., 2004), the variance in the flooding of wetlands supplied by river pulses will also likely increase. This simulation investigates the effect of a 50% increase in river stage variance on wetland development. This variance could be achieved by either raising the level of the inflow weir (by 0.3048 m in this case), or by changing the variance in the river stage itself by the same amount. Both approaches produced the same results shown in this simulation. After 100 years of development, the wetland had a similar open water area as the baseline simulation, but

158

the mudflat expanded to most of the area previously covered by marsh (Fig. 5.7b). The marsh area in this scenario became limited to a narrow fringing band along the wetlands edge.

5.5.1.3 Simulation 3 - pulsing removed, 2005 hydroperiod

In 2005 the wetland received a steady flow of pumped river water that was roughly equal in volume to the amount delivered by natural flood pulses in 2004 (See

Chapter 3). To see what the potential long-term effects of such a hydroperiod would be, a 100-year extrapolation of 2005 steady-flow hydroperiod was conducted (Fig. 5.7c). In this simulation the wetland loses the mudflat habitat that was present in the baseline simulation. The model also shows that under a steady-flow hydraulic regime the total wetland macrophyte productivity is about the same as during pulsing conditions. This result is in contrast to what was reported for this site in Chapter 4 (Fig. 4.5) where it was observed that the net macrophyte production decreased during steady-flow conditions.

This contradiction is an artifact of the grouped functional community used in the macrophyte growth model (See section 5.5.5 Model limitations, for a detailed discussion of this model limitation). Total potential macrophyte productivity was calibrated to

Typha growth because it is the dominant species in the wetland (Chapter 4; Table 4.3).

The productivity of Typha in this wetland, however, is much greater than the other emergent macrophytes in the wetland (Chapter 4; Table 4.4), but the model does not separate among plant species. Therefore macrophyte production may be overestimated by the model in the higher elevation edge areas (e.g. 725.00 ft MSL and above).

159

5.5.1.4 Simulation 4 –sediment accretion

Sedimentary processes within wetlands are intimately connected with many wetland functions including water quality and morphological changes to the wetland basin itself. Much of the existing sedimentation research comes from coastal work, with only a much smaller proportion focused on freshwater wetlands. More in-depth knowledge of sedimentary processes in freshwater wetlands is needed not only for answering questions of basic wetland science, but also for practical applications and wetland management. Understanding the long-term effects of sedimentation processes can be used to improve wetland management and design and refine predictions about the functional lifetime of wetland systems (Harter and Mitsch, 2003). The long-term effect of sediment accretion was simulated using deposition data collected in the two created emergent marshes adjacent to the created riparian oxbow modeled in this study. In a study by Harter and Mitsch (2003), the rate of sediment deposition ranged from

1.82 cm yr-1 to 9.23 cm yr-1 as measured with horizon markers. These numbers are likely high estimates as some of the sites measured were eroded, suggesting that sedimentary processes are not uniform throughout a wetland. In the unit model presented here, sedimentation is modeled as being uniform across the wetland basin, so deposition from the low end (1.82 cm yr–1) of the estimates provided by Harter and Mitsch (2003) were used in the simulation. After 100 years, the effects of sedimentation on wetland succession were most pronounced in the lower elevation contours. The 723.00 and

723.43 ft MSL contours that in the previous simulations were openwater areas filled in enough to become covered with emergent macrophytes (Fig. 5.7d). This transition from openwater to mudflat to macrophytes occurs between the 4th and 7th decade of

160

successional development (Fig. 5.8a,b). The rest of the wetland in the simulation became dominated by woody vegetation. With the 724.10 and 724.25 ft MSL contours transitioning to woody vegetation in the 6th and 7th decades (Fig. 5.8c,d). The elevations higher than 725.00 ft MSL were not significantly affected by the processes of sediment accretion (Fig. 5.8e,f,g). If the mean sedimentation rate of 4.9 cm yr-1 reported by Harter and Mitsch (2003) is used instead of the lower bound estimate, then the open water and marsh areas convert to marsh and forested areas respectively in half the time of the presented simulation.

5.5.1.5 Simulation 5 – beaver ponding

Other studies have investigated the impacts of “ecosystem engineer” species (e.g.

Jones et al., 1994, 1997; Alper, 1998; Mitsch and Jørgensen, 2004) on wetland hydroperiod and wetland plants (Sturtevant, 1998). In our model we have simulated the effect of beaver impounding by raising the outflow weir 0.3048 m. The model predicts that wetland will become a static edged pond with a narrow fringe of emergent macrophytes bounded by a ring of trees (Fig. 5.7e).

5.5.1.6 Simulation 6 – 10 years of Olentangy River data.

5.5.1.6.1 Ecosystem succession

The past 10 years of river stages in the Olentangy River were extrapolated over

100 years to provide a realistic simulation for the successional development of the created riparian wetland (Fig. 5.9). Over a 100-year period, the simulation shows that the lowest contour remains an area of open water with no plant colonization (Fig. 5.9a). In

161

drought years, the model predicts that the second lowest contour, 723.43 ft MSL, will be temporarily colonized with a small amount of biomass, but that this will not last when wet conditions return (Fig. 5.9b). The model also suggests that the “mudflat” contour from the baseline simulation will be colonized with marsh vegetation in all but the wettest years (Fig. 5.9c). This indicates that the reason the oxbow currently has not seen emergent macrophytes growing in this part of the wetland is because propagules have not reached this location and not because conditions are unfavorable for germination. In the

724.25 ft MSL contour where the baseline simulation predicts marsh vegetation, simulation 6 indicates the potential for the ecosystem to switch to forested wetland conditions (Fig. 5.9d). The 30-year span of woody coverage has the potential to be initiated and ended in each decadal water level cycle. The forested wetland submodel has a random subroutine (Fig. 5.4) that allows for the observation that trees that are adapted to wetland conditions sometimes occasionally survive prolonged inundated conditions.

This simulation underscores Neiring’s (1989) contention that chance and coincidence can play important roles in wetland ecosystem development. In this case, the chance event is forested wetland plants surviving a flood, allowing them to become an established community, and later succumbing to a flood of similar magnitude (Fig. 5.9d). Thus chance can have intermediate-term impacts on wetland succession as allogenic forcing functions can cause a reversal in ecosystem state to an earlier successional stage. Higher contours (Fig 5.9e,f,g) show little variance over the course of the simulation, indicating that flooding at these elevations is rare and not a significant stress to the ecosystem.

162

5.5.1.6.2 Total productivity

The model output was then analyzed after 8 simulation years (to correspond with

the wetlands current age) and after 100 years (to determine a potential long term end

point for the wetland’s development. At 8 years, the wetland was predicted to have a

total macrophyte productivity of 15,580 kg and a total basal area of woody plants of

10 m2 (Table 5.7). The model over estimates emergent macrophytes production by 51%

compared to the 10,000 kg value reported in Chapter 4 (Fig. 4.5). The overestimation of

macrophyte production during the eighth year occurred because the model does not

discriminate between deepwater emergent (e.g Typha spp.) and littoral species (e.g.

Juncus or Carex species). If the model output at the highest elevations ( > 725.00 ft

MSL) is corrected to reflect the lower potential peak production of these littoral species rather than Typha (Chapter 4; Fig. 4.5), then the total estimated production drops to

11,500 kg, which is only 15% different from what was actually measured. The areal

expanse of the predicted vegetation communities, however, is accurate at 8 years. The

drop in macrophyte production in the 724.43 ft MSL contour at the 100 year mark was

caused by the effect of the woody vegetation lowering the water table and making that

habitat less ideal for emergent macrophytes to grow. The model under-predicts woody

vegetation at the 8-year mark as it does not model the more rapidly growing, but

ultimately displaced, willow species that currently reside in the outflow area of the

created wetland. At 100 years, the model predicts that 15.5% of the wetland area will be

forested.

163

5.5.2 Wetland succession

Traditional successional concepts developed for upland habitats have limited

usefulness when applied to wetland dynamics. Wetlands typically remain wet over time,

exhibiting a wetland aspect rather than succeeding to upland vegetation as predicted by

hydrarch succession. Changes that occur may not necessarily be directed or orderly and

are not often predictable on the long term. Fluctuating hydrologic conditions are the

major factor controlling vegetation patterns and cyclic changes should be expected as

water levels fluctuate. Catastrophic events such as floods and droughts also play a

significant role in modifying and perpetuating these systems (Niering, 1989).

The attributes of a mature ecosystem place it in dynamic equilibrium with its

environment, and although individual species may come and go, a mature ecosystem is stable is the sense that is has built-in mechanisms that resist short –term fluctuations in the surrounding environment. Natural processes pulse regularly and the mature ecosystem responds in a pulsing steady state. Autogenic processes affect what species are able to colonize an exposed niche in a wetland ecosystem (van der Valk, 1981). Once species are established, autogenic processes replace allogenic processes as early colonists secure space and resources inhibiting the invasion of new species (Connel and Slatyer,

1977). This principle is exemplified, in the absence of additional disturbance, by the dominance of woody vegetation over macrophytes in our model.

The assumptions made by our model that propagules are evenly distributed throughout the wetland and that the functional guilds modeled are perennials are

164

reasonable. A characteristic of annuals is extended propagule longevity compared to

perennials (DeBerry and Perry, 2004). Further, anoxic conditions in submerged substrates can extend the physiological viability of seeds (Leck, 1989). Submerged habitats are commonly visited by wading birds that gather and transport seeds on mud- encrusted feathers and legs (Fenner, 1985). Our model suggests that short-term changes

in vegetation caused by periodic hydrologic pulses, are mediated by wetlands that have a

substantial, well-established seed bank. Wilcox (2004) observed this same phenomenon

in Great Lakes wetlands. Thus wetlands that are connected to rivers can be considered to

display pulse stability (Odum et al., 1995).

In wetlands, autogenic and allogenic mechanisms both contribute to vegetation

change in specific settings. The key role of water , however, makes the allogenic model

more prevalent in pulsing wetland systems (Wilcox, 2004). Individual plant species and

communities of species have affinities and physiological adaptations for certain ranges of

water depth. Water level dynamics result in shifting patterns of vegetation coverage

types. In general, deep water kills trees and emergent marsh vegetation and drawdowns

permit plant colonization. Our model supports Bedford’s (1996) claim that development and persistence of wetland ecosystems are functions of longer-term hydrologic patterns.

In simulation 6 annual changes in hydroperiod affected marsh coverage (Fig. 5.9a,b,c) had little effect on the forested wetland coverage (Fig. 5.9e,f,g). Because perennial

species, especially long-lived ones like trees, are comparatively tolerant of allogenic

disturbances, these wetlands obtain a state of pulse stability (Odum et al., 1995; Atkinson

et al., 2005).

165

The sedimentation simulation (simulation 4) predicts a typical hydrarch

succession. Although the wetland in this simulation is exposed to seasonal and annual

fluctuations in water level, these fluctuations do not allow time for the succession of plant communities to proceed. Rather it is the decadal changes associated with sedimentation that drive succession in this simulation. This model of succession is similar to one of the

succession scenarios described by Wilcox (2004) for wetlands fringing the Great Lakes.

5.5.3 Sensitivity Analysis

A detailed sensitivity analysis was performed. Initial conditions, forcing

functions, parameters, and equation/limiting factors were changed by ± 10% to observe

the corresponding changes on the state variable of greatest interest in the model. A state

variable with a larger S(p), suggests that it is more likely to change when parameters

change. Wetland depth was found to be the most sensitive of the state variables, with the

level of the outflow weir being the most influential parameter on wetland depth followed

closely by the stage of the Olentangy River (Table 5.8). Direct precipitation into the

wetland was surprisingly the least influential of the hydrologic parameters. The least

sensitive state variable was the macrophyte biomass. The most influential parameter in

this submodel was the C2 response to water depth growth coefficient and the least

influential was the Mb response to water mortality coefficient (Table 5.8). The most

influential parameters in the woody vegetation submodel were the species-specific

growth coefficients and the maximum known dimensions of the trees (Table 5.8).

166

The least influential variables were the water factor and the depth of the water table. As long as the water was deeper than the species-specific threshold, it did not have a large effect on the overall tree growth.

5.5.4 Model accuracy

Simulated water depth from the calibrated hydrologic submodel had a reasonable fit to actual measured water depths within the created riparian wetland being studied (Fig.

5.10). The mean variance between the predicted depths using 2004 data (the year the model was calibrated to) and the measured depths was 13% (equivalent to ± 7.0 cm) and had a correlation of r2 = 0.89. The hydrologic model was then validated by comparing predicted water depths for 2003 and 2005 with actual field measurements from those years. The 2003 simulation was the least accurate, with a variance of 30% (equivalent to

2 ± 15 cm) and correlation of r = 0.64. This was due to the model over estimating outflow following late summer pulses. The model tended to make this over estimation because it did not take into account the time it takes for water to move from the inflow to the outflow of the wetland. To correct this flaw, the model could be modeled as a series of cells with advective flow between the cells or by putting a delay on changes in the outflow rate equal to the current mean retention time (HRT) of the wetland. This overestimation of outflow rate was not a problem with the 2005 data (which was a year of steady-flow; Chapter 3). The 2005 hydroperiod simulation had a variance of only 5.4%

2 (equivalent to ± 2.5 cm) and correlation of r = 0.65.

167

5.5.5 Model limitations

A three-month time step of one season limited the precession of the model. A time step of a season causes the model to lose sensitivity to flood pulses. Most flood pulses in this wetland last only 10-14 days (Fink and Mitsch, in press). The model captures these pulses as an increase in the seasonal average depth in the wetland.

In reality, however, the water from pulses is often only too deep to block germination for a few weeks or for a month as opposed to for a full season. Therefore a monthly time step may be necessary.

Primary productivity of macrophytes and woody plants is a major input of biomass and hence organic matter into wetland ecosystems. Studies of freshwater marshes have shown that below-ground biomass could account for as much as two-thirds of the total biomass (Kvet and Husak, 1978; Han, 1985) but there have been few studies performed on the dynamics of underground growth (Whingham and Bailey, 1978; Mitsch and Gosselink, 2000). This is particularly true for new or constructed wetlands

(Edwards, 1992). In this model because we are primarily concerned with the potential areal coverage of vegetation types within a wetland basin, only above-ground biomass of plants was included.

The model occasionally overestimated macrophyte production (Sections 5.4.1.3 and 5.4.1.6.2) because the model does not discriminate between deepwater emergent (e.g

Typha spp.) and littoral species (e.g. Juncus or Carex species). The vegetation predicted to colonize the higher elevations (725.00 ft MSL and higher) in reality is likely to be mixed littoral macrophytes (Chapter 4; Fig. 4.3) as opposed to Typha. If the model

168

output at these elevations is corrected to reflect the lower potential peak production of these species rather than Typha, then the total estimated production becomes much more reasonable. This overestimation is only a problem in simulations where macrophyte production at the higher elevations in the wetland (7.2500 ft MSL and higher) is an issue.

The model had no provisions for the effects of competition, crowding, or standing dead. So while woody vegetation can shade out emergent macrophytes in the same plot, trees in one plot will not compete with macrophytes or trees in neighboring plots for space or water resources. Furthermore, if a tree dies from water stress, the unit is immediately open the following year for colonization in the spring season. If the unit model is to be truly expanded to a spatial scale, these issues need to be addressed.

5.6 Conclusions

This study investigated the effects of varying hydrologic inputs on the development of woody and herbaceous vegetation in a created oxbow wetland. The model was successfully able to use short-term hydrologic data to predict potential long- term ecosystem states, making it a useful tool for wetland managers. The model also showed that sedimentation processes are able to have a significant impact on the long- term development of a wetland. In order to maximize the growth of woody vegetation, the water level should be kept shallow with as little spring pulsing as possible. However, lowering the water level and eliminating pulsing would drastically reduce the habitat available for herbaceous macrophytes and would reduce both the total wetland area and the mudflat area that is currently utilized by wildlife species (Chapter 3). To optimize for wetland area, herbaceous macrophyte growth, tree growth, and wildlife utility, the water level must be allowed to fluctuate within an intermediate range (0.4-0.06 m).

169

Furthermore, even if the annual average depth is suitable for tree growth, if the range of fluctuation is too great, then woody and/or herbaceous vegetation will not be able to establish during the spring and there will be reduced vegetation cover in the wetland basin.

5.6 Acknowledgements

Support provided by U.S. Department of Agriculture NRI CSREES Award 2003-

35102-13518 and a Payne grant from the Ohio Agricultural Research and Development

Center of The Ohio State University. We also thank Dr. William Mitsch, Dr. Sven

Jørgensen, Dr. Tim Granta, Dr. Jay Martin, Dr. Robert Sykes, and Dr. Doug Alsdorf for teaching me the princiuples of ecological modeling.

170

5.7 Literature cited

Alper, J. 1998. Ecosystem “engineers” shape habitats for other species. Science 280:1195-1196.

Atkinson, R.B., J.E. Perry, and J. Cairns, Jr. 2005. Vegetation communities of 20-year- old created depressional wetlands. Wetlands Ecology and Management 13:469-478.

Baker, D.B., R.P. Richards, T.T. Loftus, J.W. Kramer. 2004. A new flashiness: Characteristics and applications to Midwestern rivers and streams. Journal of the American Water Resources Association 40:503-522.

Baldwin, A.H., K.L. McKee, I.A. Mendelssohn. 1996. The influence of vegetation, salinity, and inundation on seed banks of oligohaline coastal marshes. American journal of Botany 83:470-479.

Bedford, B.L. 1996. The need to define hydrologic equivalence at the landscape scale for freshwater wetland mitigation. Ecological Applications 6:57-68.

Bedinger, M.S. 1971. Forest species as indicators of flooding in the lower White River , Arkansas. U.S. Geological Survey Professional Paper. 750-C:248-253.

Bellport, B.P. and G.E. Burnett (eds.). 1984. Water Measurement Manual, U.S. Department of the Interior, Bureau of Reclamation, Denver, Colorado.

Benyon, R.G., S. Theiveyanathan, and T.M. Doody. 2006. Impacts of tree plantations on groundwater in south-eastern Australia. Australian Journal of Botany 54:181-192.

Botkin, D.B. 1993. Forest Dynamics: An Ecological Model. Oxford University Press, New York, 300 pp.

Boutin, C. and P.A. Keddy. 1993. A functional classification of wetland plants. Journal of Vegetation Science 4:591-600.

Brown, M.T. 2004. A picture is worth a thousand words: energy systems language and simulation. Ecological Modelling 178:83-100.

Chahine, M.T. 1992. The hydrological cycle and its influence on climate. Nature 359:373-380.

Chow, V.T. (ed). 1964. Handbook of Applied Hydrology, McGraw-Hill, New York, 1453 pp.

Connel, J.H. and R.O. Slatyer. 1977. Mechanisms of succession in natural communities and their role in community stability and organization. American Naturalist 111:1119-1144.

171

Costanza, R. 1987. Simulation modelling on the Macintosh using STELLA. BioScience 37:129-132.

Costanza, R., and T. Maxwell. 1991. Spatial ecosystem modelling using parallel processors. Ecological Modelling 58:159-183.

Cowardin, L.M., V. Carter, F.C. Golet, E.T. LaRoe. 1979. Classification of Wetlands and Deepwater Habitats of the United States, 103 pp. Washington DC. U.S. Department of the Interior, U.S. Fish and Wildlife Service.

Dawson, T.E. 1996. Determining water use by trees and forests from isotopic, energy balance, and transpiration analysis: The roles of tree size and hydraulic lift. Tree Physiology 16:263-272.

DeBerry, D.A. and J.E. Perry. 2004. Primary succession in a created freshwater wetland. Castanea 69:185-193. de Rosney, P. and J. Polcher. 1998. Modelling root water uptake in a complex land surface scheme coupled to a GCM. Hydrologic Earth Systems Science 2:239-255.

Edwards, G.S. 1992. Root distribution of soft-stem bulrush (Scirpus validus) in a constructed wetland. Ecological Engineering 1:239–243.

Ellison, A.M. and B.L Bedford. 1995. Response of a wetland vascular plant community to disturbance: a simulation study. Ecological Applications 5:109-123.

Feddes, R.A., H. Hoff, M. Bruen, T. Dawson, P. de Rosnay, P. Dirmeyer, R. Jackson, P. Kabat, A. Kleidon, A. Lilly, and A.J. Pitman. 2001. Modelling root water uptake in hydrological and climate models. Bulletin of the American Meteorlogical Society 82:2797-2809.

Fenner, M. 1985. Seed Ecology. Chapman and Hall. London, United Kingdom.

Fink, D.F. and W.J. Mitsch. in press. Hydrology and nutrient biogeochemistry in a created river diversion oxbow wetland. Ecological Engineering.

Fitz, H.C., E.B. DeBellevue, R.Costanza, R. Boumans, T. Maxwell, L. Wainger, F.H. Sklar. 1996. Development of a general ecosystem model for a range of scales and ecosystems. Ecological Modelling 88:263-295.

Gamble, D.L. 2006. Tree growth and hydrologic patterns in forested mitigation wetlands. Masters Thesis, School of Natural Resources. The Ohio State University, Coloumbus, OH.

172

Gill, C.J. 1970. The flooding tolerance of woody species - a review. Forestry Abstract 31:671-688.

Grace, J.B and R.G. Wetzel. 1981. Phenotypic and genotypic components of growth and reproduction in Typha latifolia: experimental studies in marshes for different succession maturity. Ecology 62:789-801.

Grace, J.B. 1987. The impact of preemption on the zonation of two Typha species along lakeshores. Ecological Monographs 57:283-303.

Grace, J.B. 1989. Effects of water depth on Typha latifolia and Typha domingensis. American Journal of Botany 76:762-768.

Han, J.S. 1985. Net primary production in a marsh. The Michigan Botanist 24:55-62.

Harter, S.K. and W.J. Mitsch. 2003. Patterns of short-term sedimentation in a freshwater created marsh. Journal of Environmental Quality 32:325-334.

Hook, D.D. and C.L. Brown. 1973. Root adaptations and relative flood tolerance of five hardwood species. Forest Science 19:225-229.

Jackson, R.B, H.A. Mooney, and E.D. Shulze. 2000a. Below-ground consequences of vegetation change and their treatment in models. Ecological Applications 10:470-483.

Jackson, R.B., J.S. Sperry, and T.E. Dawson. 2000b. Root water uptake and transport: Using physiological processes in global predictions. Trends in Plant Science 5:482-488.

Jaworski, E., C.N. Raphael, P.J. Mansfield, and B.B. Williamson. 1979. Impact of Great Lakes water levels on coastal wetlands. Department of Geography-Geology, Eastern Michigan University, Ypsilanti, Michigan, USA.

Jenkins, J., Chojnacky, D., Heath, L., Birdsey, R. 2003. National-scale biomass estimators for United States tree species. Forest Science 49:12-35.

Jenkins, J.C., D.C. Chojnacky, L.S. Heath, R.A. Birdsey. 2004. Comprehensive database of diameter-based biomass regressions for North American tree species. Gen. Tech. Rep. NE-319. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station.

Johnson, W.C. 1994. Woodland expansion in the Platte River, Nebraska; patterns and causes. Ecological Monographs 64:45-84.

Jones, C.G., J.H. Lawton, M. Shchak. 1994. Organisms as ecosystem enginners. Oikos 69:373-386.

173

Jones, C.G., J.H. Lawton, M. Shchak.. 1997. Positive and negative effects of organisms as physical ecosystem engineers. Ecology 78:1946-1957.

Jørgensen, S.E., and G. Bendoricchio. 2001. Fundementals of Ecological Modelling. Elsevier, Amsterdam.

Karr, R.L. 1989. Effect of flooding on Green Tree Reservoirs. Pages 14-29 in Interim Technical Reports FY 1988-1989. Water Resources Research Institute, Mississippi Sate University, Mississippi State, MS, USA.

Keddy, P.A. and A.A. Reznicek. 1986. Great Lakes vegetation dynamics: the role of fluctuating water levels and buried seeds. Journal of Great Lakes Research 12:25-36.

Keeland, B.D. and R.R. Shartz. 1997. The effects of water-level fluctuations on weekly tree growth in a southeastern USA swamp. American Journal of Botany 84:131-139.

Keely, J.E. 1979. Population differentiation along a flood frequency gradient: physiological adaptations to flooding in Nyssa sylvatica. Ecological Monographs 49: 89-108.

Kozlowski, T.T. 1984. Plant responses to flooding of soil. BioScience 34:162-167.

Kozlowski, T.T., P.J. Kramer, and S.G. Pallardy. 1991. The physiological ecology of woody plants. Acdemic Press, San Diego, California, USA.

Kusler, J.A. and M.E. Kentula. 1990. Executive summary. In: J.A. Kusler and M.E. Kentula (eds.), Wetland Creation and Restoration, the Status of the Science. Island Press, Washington, DC, pp. 17-25.

Kvet, J. and S. Husak. 1978. Primary data on biomass and productions estimates in typical stands of fishpond littoral plant communities. In D. Dykyjova` and J. Kvet (eds.) Pond Littoral Ecosystems. Springer-Verlag, Berlin.

Leck, M.A. 1989. Wetland seed banks. In: M.A. Leck, V.T. Parker, and R.L. Simpson (eds.), Ecology of Seed Banks. Academic Press, Inc. San Diego, California.

Malanson, G.P. 1993. Riparian landscapes. Cambridge University Press, Cambridge, UK.

Martin, J.F., E. Reyes, P. G. Kemp, H. Mashriqui, J.W. Day Jr. 2002. Landscape modelling of the Mississippi Delta. BioScience 52:357-366.

Middleton, B. 2000. Hydrochory, seed banks, and regeneration dynamics along the landscape boundaries of a forested wetland. Plant Ecology 146:169-184.

174

Mitsch, W.J. and B.C. Reeder. 1991. Modelling nutrient retention of a freshwater coastal wetland: estimating the roles of primary productivity, sedimentation, resuspension and hydrology. Ecological Modelling 54:151-187.

Mitsch, W.J and J.G. Gosselink. 2000. Wetlands, 3rd ed. John Wiley and Sons, Inc, New York.

Mitsch, W.J. and S.E. Jørgensen. 2004. Ecological Engineering and Ecosystem Restoration. John Wiley and Sons, Inc. Hoboken, NJ.

Nilsson, C. 1984. Effect of stream regulation on riparian vegetation. Pp. 93-106. In: A. Lillehammer and S.J Saltveit (eds) Regulated Rivers. Universitetforlagat A/S Oslo, Norway.

Neiring, W.A. 1989. Wetland vegetation development. In: S.K. Majumdar, R.P. Brooks, F.J. Brenner, R.W. Tiner (eds.), Wetlands Ecology and Conservation: Emphasis in Pennsylvania. Pp. 103-113. Pennsylvania Academy of Science, Easton PA.

Nilsson, C. 1992. Conservation management of riparian communities. Pp. 352-372 In: L. Hanssson, (ed) Ecological Principles of Nature Conservation. Elsevier Applied Science, London, U.K.

Nilsson, C., G. Grelsson, M. Johnson, and U. Sperens. 1988. Can rarity and diversity be predicted in vegetation along riverbanks? Biological Conservation 44:201-212.

Niswander, S.F. and W.J. Mitsch. 1995. Functional analysis of a 2-year old created in stream wetland: hydrology, phosphorus retention, and vegetation survival and growth. Wetlands 15:215-225.

Noon, K. 1996. A model of created wetland primary succession. Landscape and Urban Planning 34:97-123.

Odum, H.T. 1983. Systems Ecology: An Introduction. Wiley, New York.

Odum W.E., E.P. Odum, H.T. Odum. 1995. Nature’s pulsing paradigm. Estuaries 18:547–555.

Pearlstine, L., H. McKeller, and W. Kitchens, 1985. Modelling the impacts of river diversions on bottomland forest communities in the Santee River floodplain, South Carolina, Ecological Modelling 29:281-302.

Petrides, G.A. 1988. Eastern Trees. Houghton Mifflin Co., Boston, MA, 272 pp.

Phipps, R.L. 1979. Simulation of wetlands forest vegetation dynamics. Ecological Modelling 29:257-288.

175

Poiani, K.A. and W.C. Johnson. 1993. A spatial simulation model of hydrology and vegetation dynamics in semi-permanent prairie wetlands. Ecological Applications 3:279-293.

Richmond, B.S., S. Peterson, and P. Vescuso. 1987. An Academic User’s Guide to STELLATM. High Performance Systems, Lyme, NH, Pp. 392

Sargent, C.S., 1933. Manual of the Trees of North America. The Riverside Press, Cambridge, MA, 910 pp.

Shay, J.M. and C.T. Shay. 1986. Prairie marshes in western Canada, with specific reference to the ecology of five emergent macrophytes. Canadian Journal of Botany 64:443-454.

Squires, L. and A.G. van der Valk. 1992. Water-depth tolerances of the dominant emergent macrophytes of the Delta Marsh, Manitoba. Canadian Journal of Botany 70:1860-1867.

Sturtevant, B.R. 1988. A model of wetland vegetation dynamics in simulated beaver impoundments. Ecological Modelling 112:195-225.

Toner, M. and P. Keddy. 1997. River hydrology and riparian wetlands: a predictive model for ecological assembly. Ecological Applications 7:236-246. van der Valk, A.G., and C.B. Davis. 1978. The roles of seed banks in the vegetaion dynamics of prarie glacial marshes. Ecology 59:322-335. van der Valk, A.G. 1981. Succession in wetlands: a Gleasonian approach. Ecology 62:688-696.

Vannote, R.L., G.W. Minshall, K.W. Cummins, J.R. Sedell, and C.E. Cushing. 1980. The river continuum concept. Canadian Journal of Fisheries and Aquatic Science. 37:130-137.

Wang, N., W.J. Mitsch, S. Johnson, W.T. Acton. 1998. Early hydrology of a newly constructed riparian mitigation wetland at the Olentangy River Wetland Research Park. Pp. 247-254. In: W.J. Mitsch (ed.) Olentangy River Weltand Research park at The Ohio State University, Annual Report 1997.

Wang, N. and W.J. Mitsch. 2000. A detailed ecosystem model of phosphorus dynamics in created riparian wetlands. Ecological Modelling 126:101-130.

Welling, C.H., R.L. Pederson, and A.G van der Valk. 1988a. Recruitment from the seed bank and the development of zonation of emergent vegetation during a draw-down in a prairie wetland. Journal of Ecology 76:483-496.

176

Welling, C.H., R.L. Pederson, and A.G van der Valk. 1988b. Temporal patterns in recruitment from the seed bank during draw-downs in a prairie wetland. Journal of Applied Ecology 76:483-496.

Whigham, D.F. and S.E. Bailey. 1978. Nutrient dynamics in freshwater wetlands. In: Greeson, P.E., J.R. Clark, J.E. Clark (Eds.), Wetland functions and values: The state of our understanding. American Water Resources Association, Minnealpolis, MN. Pp. 4688-4798.

Wilcox, D.A. 2004. Implications of hydrologic variability on the succession of plants in Great Lakes wetlands. Aquatic Ecosystems Health and Management 7:223-231.

177

Figure 5.1 Hydrarch succession in wetlands with reversals of successional stage caused by allogenic disturbance.

178

OPEN MUDFLAT MARSH FORESTED WATER WETLAND

179

Figure 5.2 Odum diagram of interactions between hydraulic forcing functions and potential ecosystem states (open water, mudflat, marsh, and forested wetland). The forcing functions are precipitation, solar inputs, river stage or pumped inflow, and the seed bank in the wetland. The ecosystem state selected as a result of water depth is indicated by the order in which they are located in the Odum diagram.

180

Precipitation

Solar

Stage

River Flow Wetland Open water Water storage & Water depth Outflow Mudflat

Sediments Marsh

Forested wetland Nutrients

Seed bank and propagules

181

TM Figure 5.3 STELLA diagram of the hydrologic submodel. Inputs are river (Rstage), rainfall (Pt), and temperature (T). Output is the surface elevation of the water in the wetland (MSL) in both daily and in seasonal time steps. The hydrologic input to the model from the river can be switched from natural river flood pulsing (Rstage and

Pipeinflow subroutine) to a predetermined artificial pumping rate (Pump).

182

~ ~ HI Rainfall T

Pump Pt ET alpha ~

Qout Qin Q ~ RStage Weirout ~ MSLw PipeA GW in GW out Pipewater Weirhead

PipeD WL Pipeinflow Head

183

Figure 5.4 STELLATM diagram of the woody vegetation submodel. Inputs are the

surface elevation of the wetland water (MSLW) and the surface elevation of wetland

sediments (MSLL). Output is the diameter (TD) and height (TH) of tree production and the hydraulic lift (H) due to the trees.

184

TOD

HL Dmax

Dw WF b3

growth Dw Hmax G b2

TH

MSLL DegD

TD ~ Reset MSLw

Date Counter

Production Death TD

Mort MinG

Random number generator

185

Figure 5.5 STELLATM diagram of the macrophyte vegetation submodel. Inputs are surface elevation of the wetland water (MSLW), the surface elevation of the wetland sediments (MSLL), the diameter of and trees (TD), and the hydraulic lift (HL). Output is the biomass production of the emergent macrophytes (MacB).

186

TD HL

MSLw ~

Germ MSLL c2 WD ProbG

Herbaceous Growth Date Counter

Herbaceous Herbaceous MacB NAPP Death

Date Counter WD c3 ProbD

187

Figure 5.6 Site map indicating the land surface elevation contours (MSL ft) at which succession simulations were conducted.

188

189

Figure 5.7 Ecosystem state at the end of five different100 year simulations. Simulation results are a) 2004 hydroperiod, b) 2004 hydroperiod with the variance of the pulse peaks increased by 50 %, c) 2005 hydroperiod (steady flow), d) 2004 hydroperiod with sediment accretion, and e) 2004 hydroperiod with the outflow weir height raised by 1 ft

(0.3048 m).

190

(a) (b) (c)

(d) (e)

191

Figure 5.8 Time series of the simulated macrophyte and woody biomass in the created wetland according to a 100-year extrapolation of the stage of the Olentangy River from

April 2004 – March 2005 with a sediment deposition submodel. Each graph represents a different elevation contour (Fig. 5.6) within the wetland basin, with a) corresponding to the 723.00 ft MSL contour, b) the 723.43 ft MSL contour, c) the 724.10 ft MSL contour, d) the 724.25 ft MSL contour, e) the 725.00 ft MSL contour, f) the 726.00 ft MSL contour, and g) the 727.00 ft MSL contour.

192

(a) (b)

5.0 5000 5.0 5000 723.00 ft MSL Macrophytes 723.43 ft MSL Macrophytes 4.0 4000 4.0 4000 ) ) ) ) 723.00 ft MSL Woody vegetation -2 723.43 ft MSL Woody vegetation -2 -2 -2 3.0 3000 3.0 3000

2.0 2000 2.0 2000 biomass (kg m biomass (kg m vegetation (kg m vegetation (kg m Simulated macrophyte Simulated macrophyte 1.0 1000 1.0 1000 Siumulated biomass of woody Siumulated biomass of woody

0.0 0 0.0 0 0 102030405060708090100 0 102030405060708090100 Year of simulation (yr) Year of simulation (yr)

(c) (d)

5.0 5000 5.0 5000 724.10 ft MSL Macrophytes 724.25 ft MSL Macrophytes 4.0 4000 4.0 4000 ) ) ) ) -2 724.10 ft MSL Woody vegetation -2 -2

-2 724.25 ft MSL Woody vegetation 3.0 3000 3.0 3000

2.0 2000 2.0 2000 biomass (kg m (kg biomass biomass (kg m vegetation (kg m (kg vegetation vegetation (kg m Simulated macrophyte macrophyte Simulated 1.0 1000 Simulated macrophyte 1.0 1000 Siumulated biomass of woody of woody biomass Siumulated Siumulated biomass of woody 0.0 0 0.0 0 0 102030405060708090100 0 102030405060708090100 Year of simulation (yr) Year of simulation (yr) (e) (f)

5.0 5000 5.0 5000

725.00 ft MSL Macrophytes )

-2 726.00 ft MSL Macrophytes 4.0 4000 4.0 4000 ) ) ) -2 -2 725.00 ft MSL Woody vegetation -2 726.00 ft MSL Woody vegetation 3.0 3000 3.0 3000

2.0 2000 2.0 2000 biomass (kg m biomass (kg m vegetation (kg m Simulated macrophyte Simulated macrophyte 1.0 1000 Siumulated biomass of 1.0 1000 woody vegetation (kg m Siumulated biomass of woody 0.0 0 0.0 0 0 102030405060708090100 0 102030405060708090100 Year of simulation (yr) Year of simulation (yr) (g) 5.0 5000

727.00 ft MSL Macrophytes 4.0 4000 ) ) 727.00 ft MSL Woody vegetation -2 -2 3.0 3000

2.0 2000 biomass (kg m vegetation (kg m

Simulated macrophyte 1.0 1000 Siumulated biomass of woody

0.0 0 0 102030405060708090100 Year of simulation (yr)

193

Figure 5.9 Time series of simulated macrophyte and woody biomass in the created wetland according to a 100-year extrapolation of the last 10 years of Olentangy River

Stage (1996-2005). Each graph represents a different elevation contour (Fig. 5.6) within the wetland basin, with a) corresponding to the 723.00 ft MSL contour, b) the 723.43 ft

MSL contour, c) the 724.10 ft MSL contour, d) the 724.25 ft MSL contour, e) the 725.00 ft MSL contour, f) the 726.00 ft MSL contour, and g) the 727.00 ft MSL contour.

194

(a) (b)

5.0 5000 5.0 5000 723.00 ft MSL Macrophytes 723.43 ft MSL Macrophytes 4.0 4000 4.0 4000 ) ) ) -2 )

723.00 ft MSL Woody vegetation -2

-2 723.43 ft MSL Woody vegetation -2 3.0 3000 3.0 3000

2.0 2000 2.0 2000 biomass (kg m biomass (kg m vegetation (kg m vegetation (kg m Simulated macrophyte 1.0 1000 Simulated macrophyte 1.0 1000 Siumulated biomass of woody Siumulated biomass of woody

0.0 0 0.0 0 0 102030405060708090100 0 102030405060708090100 Year of simulation (yr) Year of simulation (yr) (c) (d)

5.0 5000 5.0 5000 724.10 ft MSL Macrophytes 724.25 ft MSL Macrophytes 4.0 4000 4.0 4000 ) ) ) -2 ) -2 724.25 ft MSL Woody vegetation 724.10 ft MSL Woody vegetation -2 -2 3.0 3000 3.0 3000

2.0 2000 2.0 2000 biomass (kg m (kg biomass biomass (kg m vegetation (kg m (kg vegetation vegetation (kg m Simulated macrophyte macrophyte Simulated 1.0 1000 Simulated macrophyte 1.0 1000 Siumulated biomass of woody of woody biomass Siumulated Siumulated biomass of woody

0.0 0 0.0 0 0 102030405060708090100 0 102030405060708090100 Year of simulation (yr) Year of simulation (yr) (e) (f)

5.0 5000 5.0 5000

725.00 ft MSL Macrophytes 726.00 ft MSL Macrophytes 4.0 4000 4.0 4000 ) ) ) ) -2 -2 726.00 ft MSL Woody vegetation -2

-2 725.00 ft MSL Woody vegetation 3.0 3000 3.0 3000

2.0 2000 2.0 2000 biomass (kg m biomass (kg m vegetation (kg m vegetation (kg m Simulated macrophyte Simulated macrophyte 1.0 1000 1.0 1000 Siumulated biomass of woody Siumulated biomass of woody 0.0 0 0.0 0 0 102030405060708090100 0 102030405060708090100 Year of simulation (yr) Year of simulation (yr) (g) 5.0 5000 727.00 ft MSL Macrophytes 4.0 4000 ) ) 727.00 ft MSL Woody vegetation -2 -2 3.0 3000

2.0 2000 biomass (kg m vegetation (kg m

Simulated macrophyte 1.0 1000 Siumulated biomass of woody

0.0 0 0 102030405060708090100 Year of simulation (yr)

195

Figure 5.10 Comparison of simulated and actual elevation of surface waters from April

2003 – March 2006. The model was calibrated for April 2004 – March 2005 and validated for April 2003 – March 2004 and for April 2005 – March 2006.

196

197

Ecosystem State Description Open water Water depth greater than 0.01 m and no vegetation biomass Mudflat Water depth less than 0.01 m and no vegetation biomass Marsh Macrophyte biomass greater than 0 and tree diameter less than 2 cm Forested wetland Tree diameter greater than 2 cm

Table 5.1 Four potential ecosystem successional stages predicted by the simulation model.

198

Symbol Name Values Source /units State Variables V Water volume in wetland m3 Field data; Wang et al., 1998 Forcing Functions Pt Direct precipitation m day-1 NOAA; field data WL 724.5; Weir type and elevation m Field data T Temperature oC NOAA; field data ET Evapotranspiration m day-1 Chow, 1964 HI Heat index oC Chow, 1964 -1 Rstage River stage m day Field data Pump Pumped inflow from river m day-1 Field data -1 Srate Annual sediment deposition rate cm yr Harter and Mitsch, 2003 Parameters and Coefficients 3 -1 Qin Surface inflow into the wetland m day Calculated 3 -1 Qout Surface outflow from the wetland m day Calculated Aw Area of wetland 2.8 ha Field data; Wang et al., 1998 -1 WO,a Outflow weir coefficient, a day Bellport and Burnett, 1984 and calibration -1 WO,b Outflow weir coefficient, b day Bellport and Burnett, 1984; calibration WD Water depth in the wetland m Field data -1 WI Inflow weir coefficient day Bellport and Burnett, 1984; calibration PipeD 0.3096; Inflow pipe diameter m Field data PipeA Area of the pipe filled with water m2 Calculated 3 -1 GWin Groundwater flow into wetland m day Field data; calibration 3 -1 GWout Groundwater flow out of wetland m day Field data; calibration

Table 5.2 Model parameters, definitions, values, and sources for the created oxbow wetland hydrologic submodel and sediment accretion sub-routine.

199

Symbol Name Values Source /units State Variables TD Tree diameter at breast height m Botkin, 1993 Forcing Functions DW MSLL-MSLW+HL; Depth to watertable m Hydrologic simulation DegD Growing degree days Botkin, 1993 Mort Tree mortality function cm Botkin, 1993 2 WF 1-0.055(DW-WD) ; Water growth factor Phipps, 1979 MSLL Elevation of contour m Field data Parameters and Coefficients b2 Tree growth coefficient Table 5.3 b3 Tree growth coefficient Table 5.3 GT Tree growth coefficient Table 5.3 C1 0.8; Hydraulic lift coefficient Ellison and Bedford, 1995; de Rosney and Polcher, 1998 U e-C1 x Dw; Hydraulic lift coefficient Ellison and Bedford, 1995; Kozlowski et al., 1991 HL U*0.16e-C1*TH/Hmax; Hydraulic lift m Benyon et al. 2006; Kozlowski et al., 1991 2 TH 137+b2TD-b3TOD ; Tree height cm Botkin, 1993 Dmax Maximum known tree diameter cm Table 5.3 Hmax Maximum known tree height cm Table 5.3 WD MSLW-MSLL-HL; Water depth m Calculated TOD Optimum depth to water table m Table 5.3 MSLW Elevation of wetland surface water m Hydrologic simulation β0 Biomass shape coefficient Jenkins et al. 2003 β1 Biomass shape coefficient Jenkins et al. 2003 BM Biomass of woody vegetation kg Jenkins et al. 2004 MinG Minimum growth before mortality; cm Ellison and Bedford, 1995; 0.0025 calibration

Table 5.3 Model parameters, definitions, values, and sources for the created oxbow

wetland tree submodel.

200

Species Dmax Hmax GT β0 β1 b2 b3 DegD TOD (cm) (cm) Fraxinus pennsylvanica 122 2591 188.6 43.1 0.15 0.98 0.9a Acer rubrum 150 3660 213.8 -2.01 2.43 47.0 0.16 0.86 0.6a Acer saccharinum 122 3960 164.8 -1.91 2.36 62.7 0.26 1.00 0.6 Quercus palustris 152 3353 224.7 -2.01 2.43 42.2 0.14 1.00 1.0 Liquidamber styracilua 152 4267 140.0 38.5 0.10 -0.23 1.0a Populus deltoides 152 3810 212.8 -2.21 2.39 48.2 0.16 0.82 0.6 Crataegus viridis 51 1067 123.8 36.6 0.36 0.95 0.8 Betula nigra 152 2743 158.4 34.2 0.11 0.17 0.8 a values taken from Pearlstine et al. (1985)

Table 5.4 Tree species growth coefficients that can be used in the tree growth model.

Coefficients were calculated according to Botkin (1993) unless otherwise noted. GT, β0,

β1, b2, b3 and DegD for a given species were calculated using age, height, and diameter

parameters, which were taken from Petrides (1988), Sargent (1933), and Jenkins et al.

(2003). Values of TOD were taken from Pearlstine et al. (1985) or calculated using from

Pretides (1988) and Sargent (1933) as marked. The species used in the model presented

here is Populus deltoides.

201

Symbol Name Values Source /units State Variables MacB Macrophyte Biomass g m-3 Ellison and Bedford, 1995 Sturtevant, 1998 Field data Forcing Functions Wd MSLW-MSLL-HL; Water depth in m Field data; wetland Hydrologic simulation Germ Germination threshold Table 5.5 PropG Proportion of maximum of macrophyte Ellison and Bedford, 1995 growth PropD Proportion of maximum of macrophyte Ellison and Bedford, 1995; death calibration Parameters and Coefficients C2 Change in growth rate with water level Table 5.5 C3 Change in death rate with water level Table 5.5 Ga Macrophyte growth coefficient Ellison and Bedford, 1995 Ma Macrophyte mortality coefficient Ellison and Bedford, 1995; calibration Mb Macrophyte mortality coefficient Ellison and Bedford, 1995; calibration

Table 5.5.Model parameters, definitions, values, and sources for the created oxbow wetland macrophyte submodel.

202

Character Description and value of function/parameter Lifespan Perennial and Annual – Model uses an annual growth cycle Proliferation Rhizomes and Seeds – Model assumes even seed distribution Germ 0.2b; Maximum depth of germination Growth rates Linear decrease with water level increase C2 0.2 C3 1 Ga 2 Mortality rates Exponential increase with water level increase Ma 0.01 Mb 46 Representative species Typha, Carex, and Juncus, species types Functional classesa Clonal dominants, interstitial perennials, ruderal annuals a Boutin and Keddy, 1993 b Grace, 1987; 1989

Table 5.6 Growth characteristics and parameters for functional emergent macrophyte

groups used in the macrophyte growth sub-model (from Ellison and Bedford, 1993;

model calibration). The functional group used in the model presented here is a

combination of groups 1 and 3 from Boutin and Keddy (1993).

203

Elevation Woody basal area cm2 m-2 NAPP macrophytes, g m-2 8 yrs 100 yrs 8 yrs 100 yrs 723.00 0 0 0 0 723.43 0 0 0 0 724.10 0 0 0 0 724.25 0 0.47 977 875 725.00 1.19 5530 875 0 726.00 1.21 5560 523 0 727.00 1.19 5520 523 0 Total production 1.00 m2 4600 m2 15100 kg 8750 kg

Table 5.7 Predicted basal area and net above-ground primary productivity (NAPP) per square meter of each elevation zone at the end of 8 (present year) and 100 years given a

100 year extrapolation of the past 10 years of Olentangy River flow. “Total production” refers to the entire wetland basin (multiplying the results from the unit model by the area of the wetland within the given contour band).

204

State Parameters

Variable

Less Sensitivity More

Wetland Pt GWin WI PipeD WO,b GWo WO,a Rstage WL

More Depth

Woody Wf Wd Mort TSD HL b2 Dmax MinG GT DegD Hmax b3 Basal Area 205 Sensitivity Macrophyte Mb Germ C3 Ma Wd Ga C2

Less Biomass

Sx = (Δx/x)/(Δp/p), Δp/p = ± 10%, and the order of absolute sensitivity are shown in the table; p = parameter, x = state variable.

Table 5.8 Comparison of the state variable sensitivities (Sx) to different model parameters.

205

REFERENCES

Ahn, C. and W.J. Mitsch. 2002. Scaling considerations of mesocosm wetlands in simulating large created freshwater marshes. Ecological Engineering 18:327-342.

Alper, J. 1998. Ecosystem “engineers” shape habitats for other species. Science 280:1195-1196.

American Public Health Association. 1998. Standard Methods for the Analysis of Wastewater, 20 ed., APHA, Washington, DC.

Anderson, C.J. and W.J. Mitsch. 2005. Effect of pulsing on macrophyte productivity and nutrient uptake: a wetland mesocosm experiment. American Midland Naturalist 154:305-319.

Atkinson, R.B., J.E. Perry, J. Cairns Jr. 2005. Vegetation communities of 20-year-old created depressional wetlands. Wetlands Ecology and Management 13:469–478.

Baker, D.B., R.P. Richards, T.T. Loftus, J.W. Kramer. 2004. A new flashiness: Characteristics and applications to Midwestern rivers and streams. Journal of the American Water Resources Association 40:503-522.

Baldwin, A.H., K.L. McKee, I.A. Mendelssohn. 1996. The influence of vegetation, salinity, and inundation on seed banks of oligohaline coastal marshes. American journal of Botany 83:470-479.

Bedford, B.L. 1996. The need to define hydrologic equivalence at the landscape scale for freshwater wetland mitigation. Ecological Applications 6:57-68.

Bedinger, M.S. 1971. Forest species as indicators of flooding in the lower White River Valley, Arkansas. U.S. Geological Survey Professional Paper. 750-C:248-253.

Bellport, B.P. and G.E. Burnett (eds.). 1984. Water Measurement Manual, U.S. Department of the Interior, Bureau of Reclamation, Denver, Colorado.

Benyon, R.G., S. Theiveyanathan, T.M. Doody. 2006. Impacts of tree plantations on groundwater in South-Eastern Australia. Australian Journal of Botany 54:181-192.

206

Blahnik, T. and J.W. Day. 2000. The effects of varied hydraulic and nutrient loading rates on water quality and hydrologic distributions in a natural forested treatment wetland. Wetlands 20:48-61.

Botkin, D.B. 1993. Forest Dynamics: An Ecological Model, Oxford University Press, New York.

Boustany, R.G., C.R. Crozier, J.M. Rybczyk, R.R. Twilley. 1997. Denitrification in a south Louisiana wetland forest receiving treated sewage effluent. Wetlands Ecology and Management 4:273-283.

Boutin, C. and P.A. Keddy. 1993. A functional classification of wetland plants. Journal of Vegetation Science 4:591-600.

Brown, M.T. 2004. A picture is worth a thousand words: energy systems language and simulation. Ecological Modelling 178:83-100.

Brown, S.L. 1981. A comparison of the structure, primary productivity, and transpiration of cypress ecosystems in Florida. Ecological Monographs 51:405-415.

Campbell, D.A., C.A. Cole, R.P. Brooks. 2002. A comparison of created and natural wetlands in Pennsylvania, USA. Wetlands Ecology and Management 10:41-49.

Carpenter, S.R. 1998. The need for large-scale experiments to assess and predict the response of ecosystems to perturbation. In: Pace, M.L. and P.M. Groffman (eds.), Successes, Limitations, and Frontiers of Ecosystem Science. Springer-Verlag, NY.

Casanova, M.T and M.A. Brock. 2000. How do depth, duration, and frequency of flooding influence the establishment of wetland plant communities. Plant Ecology 147:237-250.

Casey, R.E. and S.J. Klaine. 2001. Nutrient attenuation by a riparian wetland during natural and artificial runoff events. Journal of Environmental Quality 30:1720-1731.

Chahine, M.T. 1992. The hydrological cycle and its influence on climate. Nature 359:373-380.

Chow, V.T. (ed). 1964. Handbook of Applied Hydrology, McGraw-Hill, New York.

Cole, C.A. and D. Schafer. 2002. Section 404 wetland mitigation and permit success criteria in Pennsylvania, USA, 1986-1999. Environmental Management 30:508-515.

Connel, J.H. and R.O. Slatyer. 1977. Mechanisms of succession in natural communities and their role in community stability and organization. American Naturalist 111:1119-1144.

207

Connell, J.H. 1978. Diversity in tropical rainforest and coral reefs. Science 199:1302-1310.

Costanza, R. 1987. Simulation modeling on the Macintosh using STELLA. BioScience 37:129-132.

Costanza, R., and T. Maxwell. 1991. Spatial ecosystem modeling using parallel processors. Ecological Modelling 58:159-183.

Cowardin, L.M., V. Carter, F.C. Golet, E.T. LaRoe. 1979. Classification of Wetlands and Deepwater Habitats of the United States, 103 pp. Washington DC. U.S. Department of the Interior, U.S. Fish and Wildlife Service.

Cronk J.K. and W.J. Mitsch. 1994. Periphyton productivity on artificial and natural surfaces in four constructed freshwater marshes under different hydrologic regimes. Aquatic Botany 48:325-342.

Day, J. W., Jr., T. J. Butler, W. G. Conner. 1977. Productivity and nutrient export studies in a cypress swamp and lake system in Louisiana. In: M. Wiley, (ed.) Estuarine Processes, Vol. II. Academic Press, New York.

Day J.W., D. Pont, P.F. Hensel, C. Ibanez. 1995a. Impacts of sea-level rise on deltas in the Gulf of Mexico and the Mediterranean: the importance of pulsing to sustainability. Estuaries 18:636–647.

Day J.W., C.J. Madden, R.R. Twilley, R.F. Shaw, B.A. McKee, M.J. Dagg. 1995b. The influence of Atchafalaya River discharge on Fourleague Bay, Louisiana (USA). In: Dyer K.R. and R.J. Orth (eds), Changes in Fluxes in Estuaries. Olsen and Olsen, New York.

Day, J.W., R.R. Lane, R.F. Mach, C.G. Brantley, M.C. Daigle. 1999. Water chemistry dynamics in Lake Pontchartrain, Louisiana, during the 1997 opening of the Bonnet Carre Spillway. Proceedings of Recent Research in Coastal Louisiana. Lafayette, LA.

Day, J.W., L.D. Britsch, S. Hawes, G. Shaffer, D.J. Reed, D. Cahoon. 2000. Pattern and process of land loss in the Mississippi Delta: a spatial and temporal analysis of wetland habitat change. Estuaries 23:425-438.

Dawson, T.E. 1996. Determining water use by trees and forests from isotopic, energy balance, and transpiration analysis: The roles of tree size and hydraulic lift. Tree Physiology 16:263-272.

DeBerry, D.A. and J.E. Perry. 2004. Primary succession in a created freshwater wetland. Castanea 69:185-193.

208

de Rosney, P. and J. Polcher. 1998. Modelling root water uptake in a complex land surface scheme coupled to a GCM. Hydrologic Earth Systems Science 2:239-255.

Dubbe, D.R., E. G. Graver, and D.C. Pratt. 1998. Production of cattail (Typha spp.) biomass in Minnesota, USA. Biomass 23:93-185.

Edwards, G.S. 1992. Root distribution of soft-stem bulrush (Scirpus validus) in a constructed wetland. Ecological Engineering 1:239–243.

Ellison, A.M. and B.L Bedford. 1995. Response of a wetland vascular plant community to disturbance: a simulation study. Ecological Applications 5:109-123.

Feddes, R.A., H. Hoff, M. Bruen, T. Dawson, P. de Rosnay, P. Dirmeyer, R. Jackson, P. Kabat, A. Kleidon, A. Lilly, A.J. Pitman. 2001. Modeling root water uptake in hydrological and climate models. Bulletin of the American Meteorological Society 82:2797-2809.

Fenner, M. 1985. Seed Ecology. Chapman and Hall. London, United Kingdom.

Fennessy, M.S., C. Brueske, W.J. Mitsch. 1994. Sediment deposition patterns in restored freshwater marshes using sediment traps. Ecological Engineering 3:409-428.

Fennessy, M.S., and J. K. Cronk. 2001. Wetland Plants: Biology and Ecology. CRC Press.Boca Raton FL.

Fink, D.F. and W.J. Mitsch. 2004. Seasonal and storm event nutrient removal by a created wetland in an agricultural watershed. Ecological Engineering 23:313-325.

Fink, D.F. and W.J. Mitsch. in press. Hydrology and nutrient biogeochemistry in a created river diversion oxbow wetland. Ecological Engineering.

Fitz, H.C., E.B. DeBellevue, R. Costanza, R. Boumans, T. Maxwell, L. Wainger, F.H. Sklar. 1996. Development of a general ecosystem model for a range of scales and ecosystems. Ecological Modelling 88:263-295.

Faulkner, S.P., and C.J. Richardson. 1989. Physical and chemical characteristics of freshwater wetland soils. In: Hammer, D.A. (ed.), Constructed Wetlands for Wetland Wastewater Treatment. Lewis Publishers.

Fernandez, J.M., M.A.E. Selma, F.R. Aymerich, M.T.P. Saez, M.F.C Fructuoso. 2005. Aquatic birds as indicators of trophic changes and ecosystem deterioration in the Mar Menor lagoon (SE Spain). Hydrobiologia 550:221-235.

Fredrickson, L.H. and T.S. Taylor. 1982. Management of Seasonally Flooded Impoundments for Wildlife. Resource Publication 148. US Fish and Wildlife Service, Washington, DC.

209

Gagliano S, K. Meyer-Arendt, K. Wicker. 1981. Land loss in the Mississippi River Deltaic Plain. Transactions Gulf Coast Association of Geological Societies 31: 295–300.

Galat, D.L., L.H. Frederickson, D.D. Humburg, K.J. Bataille, J.R. Bodie, J. Dohrenwend. 1998. Flooding to restore connectivity of regulated, large-river wetlands (Lower Missouri River). BioScience 48:721–733.

Gamble, D.L. 2006. Tree growth and hydrologic patterns in forested mitigation wetlands. Masters Thesis, School of Natural Resources. The Ohio State University, Coloumbus, OH.

Gathumbi, S.M., P.J. Bohlen, D.A. Graetz. 2005. Nutrient enrichment of wetland vegetation and sediments in subtropical pastures. Soil Science Society of America Journal 69:539-548.

Gibbs, J.P., J.R. Longcore, D.G. McAuley, and J.K. Ringelman. 1991. Use of wetland habitats by selected nongame waterbirds in Maine. U.S. Fish and Wildlife Service Fish Wildlife Resources 9. 57 p.

Gill, C.J. 1970. The flooding tolerance of woody species - a review. Forestry Abstracts 31:671-688.

Giovanni, S.GT. and D.M.L. Da Motta Marques. 1998. Establishment of three emergent macrophytes under different water regimes. Water Science and Technology 40:233-240.

Grace, J.B and R.G. Wetzel. 1981. Phenotypic and genotypic components of growth and reproduction in Typha latifolia: experimental studies in marshes for different succession maturity. Ecology 62:789-801.

Grace, J.B. 1987. The impact of preemption on the zonation of two Typha species along lakeshores. Ecological Monographs 57:283-303.

Grace, J.B. 1989. Effects of water depth on Typha latifolia and Typha domingensis. American Journal of Botany 76:762-768.

Gusewell, S., U. Bollens, P. Ryser, F. Klotzli. 2003. Contrasting effects of nitrogen, phosphorus and water regime on first- and second-year growth of 16 wetland plant species. Functional Ecology 17:754-765.

Han, J.S. 1985. Net primary production in a marsh. The Michigan Botanist 24:55-62.

Harter, S.K. and W.J. Mitsch. 2003. Patterns of short-term sedimentation in a freshwater created marsh. Journal of Environmental Quality 32:325-334.

210

Hein, T., G. Heiler, D. Pennetzdorfer, P. Riedler, M. Schageri, F. Schiemer. 1999. The Danube Restoration Project: functional aspects and plantonic productivity in the floodplain system. Regulated Rivers 15:259-279.

Henry, C.P., C. Amoros, N. Roset. 2002. Restoration ecology of riverine wetlands: A 5- year post-operation survey on the Rhône River, France. Ecological Engineering 18:543-554.

Hensel P.F., J.W. Day, D. Pont, J.N. Day. 1998. Short-term sedimentation dynamics in the Rhone River Delta, France: The importance of riverine pulsing. Estuaries 2:52–65.

Hook, D.D. and C.L. Brown. 1973. Root adaptations and relative flood tolerance of five hardwood species. Forest Science 19:225-229.

Jackson, R.B, H.A. Mooney, E.D. Shulze. 2000a. Below-ground consequences of vegetation change and their treatment in models. Ecological Applications 10:470-483.

Jackson, R.B., J.S. Sperry, T.E. Dawson. 2000b. Root water uptake and transport: Using physiological processes in global predictions. Trends in Plant Science 5:482-488.

Jaworski, E., C.N. Raphael, P.J. Mansfield, B.B. Williamson. 1979. Impact of Great Lakes water levels on coastal wetlands. Department of Geography-geology, Eastern Michigan University, Ypsilanti, Michigan, USA.

Jenkins, J., Chojnacky, D., Heath, L., Birdsey, R. 2003. National-scale biomass estimators for United States tree species. Forest Science 49:12-35.

Jenkins, J.C., D.C. Chojnacky, L.S. Heath, R.A. Birdsey. 2004. Comprehensive database of diameter-based biomass regressions for North American tree species. Gen. Tech. Rep. NE-319. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station.

Johnson, W.C. 1994. Woodland expansion in the Platte River, Nebraska; patterns and causes. Ecological Monographs 64:45-84.

Jones, C.G., J.H. Lawton, M. Shchak. 1994. Organisms as ecosystem enginners. Oikos 69:373-386.

Jones, C.G., J.H. Lawton, M. Shchak.. 1997. Positive and negative effects of organisms as physical ecosystem engineers. Ecology 78:1946-1957.

Jørgensen, S.E., and G. Bendoricchio. 2001. Fundamentals of Ecological Modelling. Elsevier, Amsterdam.

211

Junk, W. J. 1999. The flood pulse concept of large rivers: Learning from the tropics. Archiv für Hydrobiologie 115: 261–280.

Jackson, R.B, H.A. Mooney, E.D. Shulze. 2000a. Below-ground consequences of vegetation change and their treatment in models. Ecological Applications 10:470-483.

Jackson, R.B., J.S. Sperry, T.E. Dawson. 2000b. Root water uptake and transport: Using physiological processes in global predictions. Trends in Plant Science 5:482-488.

Kadlec, R. H. 1994. Detention and mixing in free water wetlands. Ecological Engineering 3:1–36.

Kao, J.T., J.E. Titus, W. Zhu. 2003. Differential nitrogen and phosphorus retention by five wetland plant species. Wetlands 23:979-987.

Karr, R.L. 1989. Effect of flooding on Green Tree Reservoirs. Pages 14-29 in Interim Technical Reports FY 1988-1989. Water Resources Research Institute, Mississippi Sate University, Mississippi State, MS, USA.

Keddy, P. A., and A. A. Reznicek. 1986. Great Lakes vegetation dynamics: the role of fluctuating water levels and buried seeds. Journal of Great Lakes Research 12:25–36.

Keeland, B.D. and R.R. Shartz. 1997. The effects of water-level fluctuations on weekly tree growth in a southeastern USA swamp. American Journal of Botany 84:131-139.

Keely, J.E. 1979. Population differentiation along a flood frequency gradient: physiological adaptations to flooding in Nyssa sylvatica. Ecological Monographs 49: 89-108.

Kellogg, C.H., S.D Bridgham, S.A. Leight. 2003. Effects of water level , shade, and time on germination and growth of freshwater marsh plants along a simulated successional gradient. Journal of Ecology 91:274-282.

Kemp, W.M., J.E. Peterson, R.H. Gardner. 2001. Scale-dependence and the problem of extrapolation: Implications for experimental and natural coastal ecosystems. In: Gardner, R.H., W.M. Kemp, V.S. Kennedy, and J. Peterson (eds), Scaling Relationships in Experimental Ecology. Columbia University Press, NY.

Kentula, M.E., Brooks, R.B., Gwin, S.E., Holland, C.C., Sherman, A.D. Sifneos, J.C. 1992. An Approach To Improving Decision Making in Wetland Restoration and Creation Island Press, Washington, DC.

Knight, R. L., T. W. McKim, H. R. Kohl. 1987. Performance of a natural wetland treatment system for wastewater management. Journal of Water Pollution Control Federation 59:746–754.

212

Koch, M.S., and K.R. Reddy. 1992. Distribution of soil and plant nutrients along as a trophic gradient in the Florida Everglades. Journal of Soil Science Society of America 56:1492-1499.

Kozlowski, T.T. 1984. Plant responses to flooding of soil. BioScience 34:162-167.

Kozlowski, T.T., P.J. Kramer, S.G. Pallardy. 1991. The physiological ecology of woody plants. Acdemic Press, San Diego, California, USA.

Kozlowski, T.T. 2002. Physiological-ecological impacts of flooding on riparian forest ecosystems. Wetlands 22:550-561.

Kusler, J.A. and M.E. Kentula. 1990. Executive summary. In: J.A. Kusler and M.E. Kentula (eds.), Wetland Creation and Restoration, the Status of the Science. Island Press, Washington, DC.

Kvet, J. and S. Husak. 1978. Primary data on biomass and productions estimates in typical stands of fishpond littoral plant communities. In D. Dykyjova` and J. Kvet (eds.) Pond Littoral Ecosystems. Springer-Verlag, Berlin.

Lachet Instruments. 2000. Methods Manual. Lachet Instruments, Milwaukee, WI, USA.

Lane, R.R., J.W. Day, B. Thibodeaux. 1999. Water quality analysis of a freshwater diversion at Caernarvon, Louisiana Estuaries 22:327-336.

Lane R.L., J.W. Day, G.P. Kemp, D.K. Demcheck. 2001. The 1994 experimental opening of the Bonnet Carre Spillway to divert Mississippi River water into Lake Pontchartrain, Louisiana. Ecological Engineering 17:411–422.

Lane, R.R., J.W. Day, G.P. Kemp, B. Marx, E. Reyes. 2002. Seasonal and spatial water quality changes in the outfall plume of the Atchafalaya River, Louisiana, USA. Estuaries 25:30-42.

Lane, R.L., J.W. Day, D. Justica, E. Reyes, B. Marx, J.N. Day, and E. Hyfield. 2004. Changes in stoichiometric Si, N and P ratios of Mississippi River water diverted through coastal wetlands to the Gulf of Mexico. Estuarine, Coastal and Shelf Science 60:1-10

Lane, R.R., H.S. Mashriqui, G.P. Kemp, J.W. Day, J.N. Day, A. Hamilton. 2003. Potential nitrate removal from a river diversion into a Mississippi delta forested wetland. Ecological Engineering 20:237-249.

Leck, M.A. 1989. Wetland seed banks. In: M.A. Leck, V.T. Parker, and R.L. Simpson (eds.), Ecology of Seed Banks. Academic Press, Inc. San Diego, California.

Malanson, G.P. 1993. Riparian landscapes. Cambridge University Press, Cambridge, UK.

213

Maguire, C.E. 1985. Wetland replacement evaluation. Contract No. DACW-65-85-D- 0068. U.S. Army Corps of Engineers, Norfolk District, Virginia.

Martin, J.F., E. Reyes, P. G. Kemp, H. Mashriqui, J.W. Day Jr. 2002. Landscape modeling of the Mississippi Delta. BioScience 52:357-366.

Mason, C.F. and R.J. Bryant. 1975. Production, nutrient content, and decomposition of Phragmites communis, Trin. and Typha angustifolia L. Journal of Ecology 63:71-95.

McDougal, R.L., L.G. Goldsborough, B.J. Hann. 1997. Responses of a prairie wetland to press and pulse additions of inorganic nitrogen and phosphorus: Production by planktonic and benthic algae. Archiv für Hydrobiologie 140:145-167.

Megonigal, J. P., W. H. Conner, S. Kroeger, R. R. Sharitz. 1997. Aboveground production in southeastern floodplain forests: A test of the subsidy-stress hypothesis. Ecology 78:370-384.

Middleton, B. 2000. Hydrochory, seed banks, and regeneration dynamics along the landscape boundaries of a forested wetland. Plant Ecology 146:169-184.

Mitsch, W.J. and K.C. Ewel. 1979. Comparative biomass and growth of cypress in Florida wetlands. American Midland Naturalist 101:417-426.

Mitsch, W.J., C.L. Dorge J.W. Wiemhoff. 1979. Ecosystem dynamics and a phosphorus budget of an alluvial cypress swamp in southern Illinois. Ecology 60:1116-1124.

Mitsch, W.J. and W.G. Rust. 1984. Tree growth responses to flooding in a bottomland forest in northeastern Illinois. Forest Science 30:499-510.

Mitsch, W.J. and B.C. Reeder. 1991. Modelling nutrient retention of a freshwater coastal wetland: estimating the roles of primary productivity, sedimentation, resuspension and hydrology. Ecological Modelling 54:151-187.

Mitsch, W.J., J.R. Taylor, K.B. Benson. 1991. Estimating primary productivity of forested wetland communities in different hydrologic landscapes. Landscape Ecology 5:75-92.

Mitsch, W.J., J.K. Cronk, X. Wu, R.W. Nairn D.L. Hey. 1995. Phosphorus retention in constructed freshwater riparian marshes. Ecological Applications 5:830-845.

Mitsch, W.J. and R.F. Wilson. 1996. Improving the success of wetland creation and restoration with know-how, time, and self-design. Ecological Applications 6:77-83.

214

Mitsch, W.J., S. Johnson, M. Liptak. 1998. Planting and planting success of the new mitigation wetland at the Olentangy River Wetland Research Park in 1997. In: Mitsch, W.J. and V. Bouchard (eds). Olentangy River Wetland Research Park Annual Report 1997, Ohio State University, Columbus.

Mitsch, W.J., J.W. Day, J.W. Gilliam, P.M Groffman, D.L Hey, G.W. Randell, N. Wang. 1999. Reducing nutrient loads, especially nitrate-nitrogen, to surcace water, groundwater, and the Gulf of Mexico. Topic 5 Report for the Integrated Assessment on Hypoxia in the Gulf of Mexico. NOAA Coastal Ocean Program Decision Analysis Series No. 19. NOAA Coastal Ocean Program, Silver Spring, MD.

Mitsch, W.J., A.J. Horne, R.W. Nairn. 2000. Nitrogen and phosphorus retention in wetlands —Ecological approaches to solving excess nutrient problems. Ecological Engineering 14:1-7.

Mitsch, W.J and J.G. Gosselink. 2000. Wetlands, 3rd ed. John Wiley and Sons, NY.

Mitsch, W.J., J.W. Day, W. Gilliam, P.M. Groffman, D.L. Hey, G.W. Randall, N. Wang. 2001. Reducing nitrogen loading to the Gulf of Mexico from the Mississippi River Basin: strategies to counter a persistent ecological problem. BioScience 51:373-388.

Mitsch, W.J. and J.W. Day, Jr. 2004. Thinking big with whole ecosystem studies and ecosystem restoration—A legacy of H.T. Odum. Ecological Modelling 178:133-155.

Mitsch, W.J. and S.E. Jorgensen. 2004. Ecological Engineering and Ecosystem Restoration. John Wiley and Sons, Inc. Hoboken, NJ.

Mitsch, W.J., N. Wang, L. Zhang, R. Deal, X, Wu, A. Zuwerink. 2005a. Using ecological indicators in a whole-ecosystem wetland experiment. In: Jorgensen, S.E., F.L. Xu, R. Costanza (eds.), Handbook of Ecological Indicators for Assessment of Ecosystem Health, CRC Press, Boca Raton, FL.

Mitsch, W.J., J.W. Day, Jr., L. Zhang, R. Lane. 2005b. Nitrate-nitrogen retention by wetlands in the Mississippi River Basin. Ecological Engineering 24:267-278.

Mitsch, W.J., L. Zhang, C.J. Anderson, A.E. Altor, M.E. Hernandez. 2005c. Creating riverine wetlands: Ecological succession, nutrient retention, and pulsing effects. Ecological Engineering 25:521-527.

Mitsch, W.J. and J.W. Day, Jr. 2006. Restoration of wetlands in the Mississippi-Ohio- Missouri (MOM) River Basin: Experience and needed research. Ecological Engineering 26:55-69.

Molles M.C., C.S. Crawford, L.M. Ellis, H.M. Valett, C.N. Dahm. 1998. Managed flooding for riparian ecosystem restoration. BioScience 48:748–756.

215

Nairn, R.W. and W.J. Mitsch. 2000. Phosphorus removal in created wetland ponds receiving river overflow. Ecological Engineering 14:107-126.

National Research Council. 1992. Restoration of Aquatic Ecosystems. National Academy Press, Washington, DC.

Newman, S., J. Schuette, J.B. Grace, K. Ritchey, T. Fontaine, K.R. Reddy, M. Pietrucha. 1998. Factors influencing cattail abundance in the northern everglades. Aquatic Botany 60:265-280.

Neiring, W.A. 1989. Wetland vegetation development. Pp. 103-113. In: S.K. Majumdar, R.P. Brooks, F.J. Brenner, R.W. Tiner (eds.), Wetlands Ecology and Conservation: Emphasis in Pennsylvania. Pennsylvania Academy of Science, Easton PA.

Nilsson, C. 1984. Effect of stream regulation on riparian vegetation. Pp. 93-106. In A. Lillehammer and S.J Saltveit (eds.), Regulated Rivers. Universitetforlagat A/S Oslo, Norway.

Nilsson, C. 1992. Conservation management of riparian communities. Pp. 352-372 In L. Hanssson, (ed.) Ecological Principles of Nature Conservation. Elsevier Applied Science, London, UK.

Nilsson, C., G. Grelsson, M. Johnson, and U. Sperens. 1988. Can rarity and diversity be predicted in vegetation along riverbanks? Biological Conservation 44:201-212.

Niswander, S.F. and W.J. Mitsch. 1995. Functional analysis of a 2-year old created in stream wetland: hydrology, phosphorus retention, and vegetation survival and growth. Wetlands 15:215-225.

Nixon, S.W. 1988. Physical energy inputs and the comparative ecology of lake and marine ecosystems. Limnology and Oceanography 33:1005–1025.

Nixon, S. W., J.W. Ammerman, L.P. Atkinson, V.M. Berounsky, G. Billen, W.C. Boicourt, W.R. Boynton, T.M. Church, D.M. Ditoro, R. Elmgren, J.H. Garber, A.E. Giblin, R.A. Jahnke, N.J.P. Owens, M.E.Q. Pilson, S.P. Seitzinger. 1996. The fate of nitrogen and phosphorus at the land-sea margin of the North Atlantic Ocean. Biogeochemistry 35:141-180.

Noon, K. 1996. A model of created wetland primary succession. Landscape and Urban Planning 34:97-123.

Novitzki, R.P. 1982. Hydrology of Wisconsin Wetlands: Wisconsin Geological and Natural History Survey Information Circular 40, Madison, WI.

Odum, H.T. 1983. Systems Ecology: An Introduction. Wiley, New York.

216

Odum, E.P. 1990. Field experimental tests of ecosystem-level hypotheses. Trends in Ecological Evolution 5:204–205.

Odum, W.E., E.P. Odum, H.T. Odum. 1995. Nature’s pulsing paradigm. Estuaries 18:547–555.

Odum, E. P. 2000. Tidal marshes as outwelling/pulsing systems. In M. P. Weinstein and D. A. Kreeger (eds.), International Symposium: Concepts and Controversies in Tidal Marsh Ecology. Kluwer Academic Publishers, Dordrecht.

Olila, O.G, K.R. Reddy, D.L. Stites. 1997. Influence of draining on soil phosphorus forms and distribution in a constructed wetland. Ecological Engineering 14:107-126.

Pearlstine, L., H. McKeller, W. Kitchens, 1985. Modelling the impacts of river diversions on bottomland forest communities in the Santee River floodplain, South Carolina, Ecological Modelling 29:281-302.

Perez, B. C. and J. W. Day. 2000. Influence of Atchfalaya River discharge and winter frontal passage and flux in Four League Bay, Louisiana. Estuarine, Coastal and Shelf Science 50: 271-290.

Petrides, G.A. 1988. Eastern Trees. Houghton Mifflin Co., Boston, MA.

Phipps, R.L. 1979. Simulation of wetlands forest vegetation dynamics. Ecological Modelling 29:257-288.

Poiani, K.A. and W.C. Johnson. 1993. A spatial simulation model of hydrology and vegetation dynamics in semi-permanent prairie wetlands. Ecological Applications 3:279-293.

Poole, A.F., P.R. Stettenheim, F. Gill. 1992. The Birds of North America: Life Histories for the 21st century. (eds.) Alan F. Poole, Peter Stettenheim, and Frank Gill. American Ornithologists' Union; Washington, DC.

Rabalais, N. N., W. J. Wiseman, R. E. Turner, B. K. Sengupta, Q. Dortch. 1996. Nutrient changes in the Mississippi River and system responses on the adjacent continental shelf. Estuaries 19:386-407.

Rabalais, N. N., R. E. Turner, W. J. Wiseman, Q. Dortch. 1998. Consequences of the 1993 Mississippi River flood in the Gulf of Mexico. Regulated Rivers 14:161-177.

Rabalais, N.N., R.E. Turner, D. Justic, Q. Dortch, W.J. Wiseman. 1999. Topic 1 Report for the Integrated Assessment on Hypoxia in the Gulf of Mexico. NOAA Coastal Ocean Program Decision Analysis NO. 15; NOAA Coastal Ocean Program, Silver Spring, MD.

217

Rabalais, N.N., R.E. Turner, D, Scavia. 2002. Beyond science and into policy: Gulf of mexico hypoxia and the Mississippi River. BioScience 52:129-142.

Randell, G.W., D.R. Huggins, M.P. Russelle, D.J. Fuchs, W.W. Nelson, J.L Anderson. 1997. Nitrate losses through subsurface tile drainage in CRP, alfalfa and row crop systems. Journal of Environmental Quality 26:1240-1247.

Raisen, G.W. and D.S. Mitchel. 1995. The use of wetlands for the control of non-point source pollution. Water Science and Technology 32:177-186.

Raisen, G.W., D.S. Mitchel, R.L. Croome. 1997. The effectiveness of a small constructed wetland in meliorating diffuse nutrient loadings from an Australian rural catchment. Ecological Engineering 9:19-36.

Reddy, K.R. and D.A. Graetz. 1988. Carbon and nitrogen dynamics in wetland soils. Pp. 307-318. In: D.D. Hook, W.H. McKee, Jr., H.K. Smith, J. Gregory, V.G. Burrel, M.R. DeVoe, R.E. Sojka, S. Gilbert, R.Banks, L.G. Stolzy, C. Brooks, T.D. Mathews, and T.H Shear (eds.) The Ecology and Management of Wetland, Vol. 1: The Ecology of Wetlands. Timber Press, Portland, OR.

Reed, P.B., Jr. 1998. National list of plant species that occur in wetlands: Northeast (Region I). U.S. Fish and Wildlife Service, Washington, DC, Biological Report 88 (26.1).

Reinartz, J.A. and E.L. Warne. 1993. Development of vegetation in small created wetlands in southeastern Wisconsin. Wetlands 13:153-164.

Reyes E, M.L. White, J.F. Martin, G.P. Kemp, J.W. Day, A. Aravamuthan. 2000. Landscape modeling of coastal habitat change in the Mississippi Delta. Ecology 81:2331–2349.

Richardson, C.J., Nichols, D.S., 1985. Ecological analysis of wastewater management criteria in wetland ecosystems, Ecological considerations pp. 351-391. In: E.R.K. Paul J. Godfrey, Sheila Pelczarski (eds.), Wetlands Treatment of Municipal Wastewaters, Van Nostrand Reinhold Company, N/Y.

Richmond, B.S., S. Peterson, P. Vescuso. 1987. An Academic User’s Guide to STELLATM. High Performance Systems, Lyme, NH.

Rutschke, E., 1987. Waterfowl as bio-indicators. In: Diamond, A.W., F.L. Filion (eds), The value of birds. ICBP Technical Publication No. 6. 167–172.

Sargent, C.S. 1933. Manual of the Trees of North America. The Riverside Press, Cambridge, MA.

218

Shay, J.M. and C.T. Shay. 1986. Prairie marshes in western Canada, with specific reference to the ecology of five emergent macrophytes. Canadian Journal of Botany 64:443-454.

Spieles, D.J. and W.J. Mitsch. 2000. The effects of season and hydrologic and chemical loading on nitrate retention in constructed wetlands: a comparison of low- and high- nutrient riverine systems. Ecological Engineering 14:77-91.

Spink, A., R.E. Sparks, M. van Oorschot, T.W. Verhoenven. 1998. Nutrient dynamics of large river floodplains. Regulated Rivers: Research and Management 14:203-216.

Squires, L. and A.G. van der Valk. 1992. Water-depth tolerances of the dominant emergent macrophytes of the Delta Marsh, Manitoba. Canadian Journal of Botany 70:1860-1867.

Steen, D.A., J.P. Gibbs, T.A. Timmermans. 2006. Assessing the sensitivity of bird communities to hydrologic change in the eastern Great Lakes region. Wetlands 26:605-611.

Sturtevant, B.R. 1988. A model of wetland vegetation dynamics in simulated beaver impoundments. Ecological Modelling 112:195-225.

Taft, O.W., M.A. Colwell, C.R. Isola, R.J. Safran. 2002. Waterbird responses to experimental drawdown: implications for the multispecies management of wetland mosaics. Journal of Applied Ecology 39:987-1001.

Tanner, C.C., J. D’Eugenio, G.B. McBride, J.P.S. Sukias, K. Thompson. 1999. Effect of water level fluctuation on nitrogen removal from constructed wetland mesocosms. Ecological Engineering 12:67-92.

Tockner, K., F. Malard, J.V. Ward. 2000. An extension of the flood pulse concept. Hydrologic Processes 14:2861-2883.

Toner, M. and P. Keddy. 1997. River hydrology and riparian wetlands: A predictive model for ecological assembly. Ecological Applications 7:236-246.

Toth L.A., S.L. Melvin, D.A. Arrington, J. Chamberlain. 1998. Hydrologic manipulations of the channelized Kissimmee River. BioScience 48:757–765.

Townsend, C.R. 1996. Concepts in river ecology: pattern and process in the catchment hierarchy. Archiv fur Hydrobiologie Supplement 113, Large Rivers 10: 3-21.

Turner, R. E.; Rabalais, N. N., Swenson, E. M., Kasprzak, VI Romaire, T. Marine. 2005. Summer hypoxia in the northern Gulf of Mexico and its prediction from 1978 to 1995. Environmental Research 59:65-77.

219

Tuttle, C.L. and W.J. Mitsch. in review. Aquatic metabolism as an indicator of the ecological effects of hydrologic pulsing in flow-through wetlands. Ecological Indicators.

United States Bureau of Reclamation (1997). Water Measurement Manual 3ed. U.S. Department of the Interior, Bureau of Reclamation. Washington DC.

U.S. Environmental Protection Agency. 1983. Methods for Chemical Analysis of Water and Wastes. 600/4-79-020, U.S. Environmental Protection Agency, Cincinnati, OH. van der Valk, A.G. and C.B. Davis. 1978. The role of seed banks in the vegetation dynamics of prairie glacial marshes. Ecology 59:322-335. van der Valk, A.G. 1981. Succession in wetlands: a Gleasonian approach. Ecology 62:688-696.

Vannote, R.L., G.W. Minshall, K.W. Cummins, J.R. Sedell, C.E. Cushing. 1980. The river continuum concept. Canadian Journal of Fisheries and Aquatic Science 37:130-137.

Wang, N., W.J. Mitsch, S. Jjohnson, W.T. Acton. 1998. Early hydrology of a newly constructed riparian mitigation wetland at the Olentangy River Wetland Research Park. Pp. 247-254. In: W.J. Mitsch (ed.) Olentangy River Weltand Research Park at The Ohio State University, Annual Report 1997.

Wang, N. and W.J. Mitsch. 2000. A detailed ecosystem model of phosphorus dynamics in created riparian wetlands. Ecological Modelling 126:101-130.

Waters, I. and J.M Shay. 1992. Effect of water depth on population parameters of Typha glauca stand. Canadian Journal of Botany 70:349-351.

Welling, C.H., R.L. Pederson, A.G van der Valk. 1988a. Recruitment from the seed bank and the development of zonation of emergent vegetation during a draw-down in a prairie wetland. Journal of Ecology 76:483-496.

Welling, C.H., R.L. Pederson, and A.G van der Valk. 1988b. Temporal patterns in recruitment from the seed bank during draw-downs in a prairie wetland. Journal of Applied Ecology 76:483-496.

Whigham, D.F. and S.E. Bailey. 1978. Nutrient dynamics in freshwater wetlands. pp. 468-478. In: Greeson, P.E., J.R. Clark, J.E. Clark (Eds.), Wetland functions and values: The state of our understanding. American Water Resources Association, Minneapolis, MN.

Wilcox, D. A. 1993. Effects of water-level regulation on wetlands of the Great Lakes. Great Lakes Wetlands 4:1-11.

220

Wilcox, D.A. and T.H. Whillans. 1999. Techniques for restoration of disturbed coastal wetlands of the Great Lakes. Wetlands 19:835-857.

Wilcox, D.A. 2004. Implications of hydrologic variability on the succession of plants in Great Lakes wetlands. Aquatic Ecosystems Health and Management 7:223-231.

Wong, T.H.F., and N.L.G. Somes. 1995. A stochastic approach to designing wetlands for stormwater pollution control. Water Science and Technology 32:145-151.

Zedler, J.B. and J.C. Callaway. 2000. Evaluating the progress of engineered tidal wetlands. Ecological Engineering 15:211-225.

221

APPRENDIX A

STELLA CODES FOR DYNAMIC SIMULATION MODELS

222

Hydrology submodel

V(t) = V(t - dt) + (Qin + GWin + Pt - Qout - GWout - ET) * dt INIT Q = 400000

Qin = Pipeinflow*43560 GWin = 0 Pt = Rainfall*2.8*2.477*43560/5

Qout = Weirout*43560

ET = IF (T>0) THEN (IF (MSLW<723.4) THEN (2.8*2.477*43560*(1/12/10/2.54)*(16/12)*((10*T/HI)^alpha)) ELSE (2.8*2.477*43560*(1/12/18/2.54)*(16/12)*((10*T/HI)^alpha))) ELSE (0)

alpha = ((0.675*HI3)-(77.1*HI2)+(17920*HI)+(492390))*10-6

Head = MAX(0,MIN((RStage-MSLW),(RStage*(0.6*(MIN(2.3,MAX(0,(RStage-723.5)))))- 723.5)))

HI = If (T>0) THEN ((T/5)1.514)ELSE(0)

-5 -6 MSLW = IF (Q<17983) THEN (723.43+Q*2.787*(10 )) ELSE (723.43+Q*4.662*(10 )- (Q2)*7.95*(10-13))

PipeA = IF(Pipewater=0)THEN(0)ELSE(IF((((Pipewater)- (PipeD/2))/(PipeD/2))>0)then(((PI*(PipeD/2)2)*(360-2*(ARCTAN((SQRT(1- (((Pipewater)-(PipeD/2))/(Pipewater/2))2))/((((PipeD)- (PipeD/2))/(PipeD/2)))))*180/PI)/360)+((Pipewater)-(PipeD/2))*((PipeD/2)2- ((Pipewater)-(PipeD/2))2)0.5)ELSE((((360-2*(3.14+ARCTAN((SQRT(1-((Pipewater- (PipeD/2))/(PipeD/2))2))/((Pipewater-(PipeD/2))/(PipeD/2))))*180/PI)/360))+(Pipewater- (PipeD/2))*((PipeD/2)2-(Pipewater-(PipeD/2))2)0.5))

PipeD = 2

Pipeinflow = (1.9+0.19)*PipeA*SQRT(Head)

Pipewater = IF (RStage-723.5<0) THEN (0) ELSE (MIN(2,RStage-723.5))

Weirhead = IF (WL

Weirout = 10.16*(Weirhead1.436)

WL = 724.5

223

Vegetation submodels

Macrophytes MacB(t) = MacB(t - dt) + (Herb_NAPP - Herb_Death) * dt INIT MacB = 0

Herb_NAPP = IF (Date_Counter=3) THEN (0) ELSE (Herb_Growth*2640)

Herb_Death = IF (Date_Counter=3) THEN (MacB) ELSE (ProbD*MacB) TD(t) = TD(t - dt) + (Production - Death) * dt INIT TD = 0

C2 = 0.2 C3 = 1

ProbD = IF ((WD>0.08) AND (Date_Counter>1)) THEN (47*WD) (47*WD) (Date_Counter*C3*0.01*e ) ELSE (C3*0.01* e )

ProbG = IF ((Date_Counter=2) AND (WD>0.08)) THEN (0) ELSE (IF (TD<1) THEN (IF (WD<0) THEN (C2) ELSE (C2*(1-2*WD))) ELSE (0))

Macrophyte_peak_biomass = IF (Date_Counter=3) THEN (MacB) ELSE (-1)

WD = 0.3048*(MAX(MSLW-HL-MSLL,0))

Herbaceous_Growth = IF (MacB>0) THEN (ProbG) ELSE (IF (Germ=1) THEN (ProbG) ELSE (0))

Death = Mort*TD

Date_Counter = delay(COUNTER(1,5),2,4)

Trees b2 = 48.2 b3 = 0.16 C1 = 0.8 DegD = 0.82 Dmax = 152 Hmax = 2*1905 GT = 212.8 DW = MSLL-MSLW+HL

Germ = IF ((MacB=0) AND (WD<0.2) AND ((Date_Counter=1))) THEN (1) ELSE (0) growth = 0.25*Nutrient_Factor*WF*DegD*GT*TD_Reset*(1- ((TD_Reset*TH)/(Dmax*Hmax)))/(274+(3*b2*TD_Reset)-(4*b3*TD_Reset))

224

Production = IF (Mort=1) THEN (0) ELSE (growth)

(-TH/Hmax) HL = IF (TH>150) THEN (U*0.16*(e )) ELSE (0)

MinG = IF (growth<0.0025) THEN (Random_number_generator) ELSE (1)

Mort = IF (MinG<0.368) THEN (1) ELSE (0)

Random_number_generator = random(0,1)

TD_Reset = IF ((TD=0) AND (Date_Counter=1) AND (MSLL-MSLW>0)) THEN (0.2) ELSE (TD)

2 TH = IF (TD_Reset=0) THEN (0) ELSE (137+b2*TD_Reset-b3*(TD_Reset ))

TOD = 0.6

(-C *(MSL -MSL ) U = e 1 L W

2 WF = IF (DW<0.0) THEN (0) ELSE (MIN(1-0.055*(DW-TOD) ,1))

Ecosystem state determination Ecosystem_state_determinator = IF ((MacB=0) AND (TD=0) AND (WD<0.02) AND (Date_Counter=2)) THEN (“Open Water”) ELSE (IF ((MacB=0) AND (TD=0) AND (WD>0.02) AND (Date_Counter=3)) THEN (“Mud Flat”) ELSE (IF ((Date_Counter=3) AND (MacB>0)) THEN (“Macrophytes”) ELSE (IF ((Date_Counter=3) AND (TD>0)) THEN (“Trees”) ELSE (0))))

225