RAPID RESPONSE TO A CATASTROPHIC FLOOD: EFFECTS ON AQUATIC
RESOURCES IN MISSOURI RIVER RESERVOIRS
BY
ANDREW KENNETH CARLSON
A thesis submitted in partial fulfillment of the requirements for the
Master of Science
Major in Wildlife and Fisheries Science
Specialization in Fisheries
South Dakota State University
2015
iii To Laura and Dan Carlson, for guidance, patience, and unwavering love
iv ACKNOWLEDGMENTS
First and foremost, I thank my parents, Laura and Dan Carlson, to whom this
thesis is dedicated. Your everlasting love has molded me into the person I am today. I
will cherish it as long as I live. What a privilege it has been to attend South Dakota State
University to pursue natural resource science, a passion you inspired. The familial
significance of Brookings inspires me every day.
I thank the numerous teachers, coaches, environmental educators, and natural
resource professionals who furnished knowledge in biology, ecology, and fisheries
science and motivated me to learn more. I would not write these words without you.
Thank you especially to Jim Christensen, Dan Kovacich, Jacob Montgomery, and Robert
Rantanen (White Bear Lake Area High School); Drs. Leonard C. Ferrington, Rebecca A.
Montgomery, James A. Perry, Andrew M. Simons, and Bruce Vondracek (University of
Minnesota); Julie Grecian, Ron Lawrenz, and Geoff Urban (Lee and Rose Warner Nature
Center); and Kacie Carlson, Dave Crawford, Will French, Jerry Johnson, Tom Landwehr,
Don Pereira, Brad Parsons, Brian Nerbonne, Jim Levitt, Joel Stiras, and Heidi Wolf
(Minnesota Department of Natural Resources).
I thank faculty advisor, Dr. Brian Graeb, for perpetual enthusiasm and sagacious
guidance throughout the duration of this project. You opened my mind to the wonders of
fisheries research and inspired me to think broadly (at 10,000 feet and beyond) about the
implications of my study. It was an honor working in your lab. I thank my project
advisors, Dr. Mark Fincel and Chris Longhenry, for providing outstanding professional
advice, helpful comments on talks and papers, and much needed levity in the office and
field. Thank you for serving as valuable mentors during my tenure at South Dakota State
v University. I also thank Drs. Katie N. Bertrand, Michael L. Brown, Steve R. Chipps,
Larry M. Gigliotti, Kent C. Jensen, Joshua D. Stafford, Nels H. Troelstrup, and Melissa
R. Wuellner, for their professional advice, analytical assistance, and pleasant
conversation throughout the duration of my research.
I am especially grateful for the wisdom and guidance of the late Drs. David W.
Willis and Robert A. Klumb. Both provided me with invaluable insight for conducting
research and navigating graduate school. Through students like myself they inspired,
their legacies will live on forever.
I thank Jacob Fernholz, Brandon Hoffman, Ryan Johnston, and Janae Oien for
their assistance in the laboratory and field. Your patience and persistence enabled pivotal
discoveries for Missouri River management. I thank Bob Hanten, Gary Knecht, Hilary
Meyer, Kyle Potter, Jason Sorensen, and Rachel Trible of the South Dakota Department
of Game, Fish and Parks and Paul Bailey of the North Dakota Game and Fish
Department. Your assistance in the laboratory and field was invaluable. I thank Dan
James of the U.S. Fish and Wildlife Service for providing data and offering analytical
advice for my research in the Lewis & Clark Delta. I thank Alan Shiller and his lab
members from the University of Southern Mississippi for providing water sampling kits
and measuring trace element concentrations in water samples. I also thank Gry Barford,
Joel Commisso, and Justin Glessner from the University of California – Davis for their
assistance with otolith microchemistry analyses.
I thank my fellow graduate students at South Dakota State University, especially
Mark Kaemingk, Laura Heironimus, Dan Dembkowski, Bradley Smith, Jeffrey Grote,
Natalie Scheibel, Jeremy Kientz, Ryann Cressey, Brandi Crider, Mandy Orth, Sarah
vi Nevison, Diane Narem, Tait Ronningen, Alex Rosburg, David Deslauriers, and Joe
Bennett. I give special thanks to those in the Graeb/Bertrand Lab Group, including CariAnn Hayer, Jason Breegeeman, William Radigan, Eli Felts, Tobias Rapp, Darrel (Jake)
Mecham, Jacob Krause, Jessica Howell, Chad Kaiser, Matt Wagner, David Schumann,
John Lorenzen, Cassidy Gerdes, Stephen Jones, Seth Fopma, Tanner Stevens, Lyntausha
Kuehl, and Joe Bennett. Your insightful comments (most notably, “a sophisticated heat
beam … called ‘a laser’”) improved my talks and strengthened the powerhouse that is our
lab group.
Finally, I thank Terri Symens, Di Drake, Dawn Van Ballegooyen, and Kate Tvedt
for their cordial assistance with project finances and travel.
Funding for this project was provided by the Federal Aid in Sport Fish
Restoration program, Project 3M3328, Study 1526, administered by the South Dakota
Department of Game, Fish and Parks, United States Fish & Wildlife Service, South
Dakota State University Department of Natural Resource Management, and South
Dakota Agricultural Experiment Station.
Author’s Note: Use of personal pronouns (i.e., “We” in place of “I”) reflects the
collaborative nature of this research project.
vii CONTENTS
LIST OF FIGURES ............................................................................................................. x LIST OF TABLES ............................................................................................................. xv ABSTRACT ..................................................................................................................... xvii CHAPTER 1. INTRODUCTION ...................................................................................... 1 River Management .......................................................................................................... 1 Opportunities and Challenges for Missouri River Management .................................... 3 Catastrophic Flooding .................................................................................................... 6 Entrainment ..................................................................................................................... 7 Sedimentation .................................................................................................................. 9 Project Overview ........................................................................................................... 10 REFERENCES .............................................................................................................. 13 CHAPTER 2: SPATIAL HETEROGENEITY IN TRACE ELEMENT CHEMISTRY IN
MISSOURI RIVER RESERVOIRS ................................................................................. 25 Abstract ......................................................................................................................... 26 Study Site ...................................................................................................................... 28 Materials and Methods .................................................................................................. 29 Results ........................................................................................................................... 29 Discussion ..................................................................................................................... 30 Acknowledgements ....................................................................................................... 32 References ..................................................................................................................... 34 Tables ............................................................................................................................ 36 Figures ........................................................................................................................... 38 CHAPTER 3: OTOLITH MICROCHEMISTRY REVEALS NATAL ORIGINS OF
WALLEYES IN MISSOURI RIVER RESERVOIRS ..................................................... 44 Abstract ......................................................................................................................... 45 Introduction ................................................................................................................... 46 Methods ......................................................................................................................... 48 Study Area.................................................................................................................. 48 Trace Element Sampling ............................................................................................ 49 Fish Sampling ............................................................................................................ 49 viii Otolith Microchemistry.............................................................................................. 50 Statistical Analysis ..................................................................................................... 54 Results ........................................................................................................................... 55 Water Chemistry ........................................................................................................ 55 Otolith Chemistry ...................................................................................................... 56 Natal Contribution ..................................................................................................... 56 Discussion ..................................................................................................................... 57 Acknowledgments ......................................................................................................... 62 References ..................................................................................................................... 64 Tables ............................................................................................................................ 72 Figures ........................................................................................................................... 75 CHAPTER 4: THE MISSOURI RIVERSCAPE: OTOLITH MICROCHEMISTRY
FACILITATES ECOSYSTEM-BASED FISHERIES MANAGEMENT IN NORTH
AMERICA’S LONGEST RIVER .................................................................................... 79 Abstract ......................................................................................................................... 80 Introduction ................................................................................................................... 81 Methods ......................................................................................................................... 83 Study Area.................................................................................................................. 83 Trace Element Sampling ............................................................................................ 83 Fish Sampling ............................................................................................................ 84 Otolith Microchemistry.............................................................................................. 85 Statistical Analysis ..................................................................................................... 88 Results ........................................................................................................................... 89 Water Chemistry ........................................................................................................ 89 Otolith Chemistry ...................................................................................................... 90 Movement................................................................................................................... 90 Entrainment ............................................................................................................... 92 Discussion ..................................................................................................................... 94 Acknowledgments ....................................................................................................... 101 References ................................................................................................................... 102 Tables .......................................................................................................................... 110 Figures ......................................................................................................................... 115 ix CHAPTER 5: EFFECTS OF CATASTROPHIC FLOODING ON A DELTA
ECOSYSTEM IN THE MISSOURI RIVER .................................................................. 122 Abstract ....................................................................................................................... 123 Introduction ................................................................................................................. 124 Methods ....................................................................................................................... 126 Study Area................................................................................................................ 126 Fish Sampling .......................................................................................................... 127 Aquatic Habitats ...................................................................................................... 127 Statistical Analysis ................................................................................................... 127 Results ......................................................................................................................... 129 Structural Indices .................................................................................................... 129 Relative Abundance by Introduction History .......................................................... 130 Relative Abundance by Trophic Guild ..................................................................... 131 Aquatic Habitats ...................................................................................................... 132 Discussion ................................................................................................................... 132 Acknowledgments ....................................................................................................... 136 References ................................................................................................................... 138 Figures ......................................................................................................................... 148 CHAPTER 6: SUMMARY, MANAGEMENT RECOMMENDATIONS, AND
RESEARCH NEEDS ...................................................................................................... 153 APPENDICES ................................................................................................................ 161 x LIST OF FIGURES
Figure 2-1. Map of South Dakota surficial geology. Geologically young glacial sediments
predominate east of the Missouri River, whereas older Triassic (Tr), Jurassic
(J), Cretaceous (C), Tertiary (T) sediments exist west of the river. M denotes
metamorphic rocks. Image obtained from South Dakota Department of
Environment & Natural Resources
(http://www.sdgs.usd.edu/geologyofsd/geosd.html). ....................................... 38
Figure 2-2. Sr:Ca and Ba:Ca ratios for tributaries (black boxes) and select
mainstem/embayment sites (gray boxes) in Missouri River reservoirs (Oahe,
Sharpe, Francis Case [FC], and Lewis & Clark [LC]) in North Dakota and
South Dakota, USA. All tributaries and a representative set of
mainstem/embayment sites sampled in July 2012 and February 2014 are
depicted. Tributaries exhibit spatial variability (i.e., north–south gradient) in
water Sr:Ca and Ba:Ca ratios, whereas mainstem/embayment sites generally
have similar water signatures. Means with the same letter are not significantly
different (ANOVA with Tukey’s HSD test; P < 0.05). ................................... 39
Figure 2-3. Mean water Sr:Ca values at 20 collection sites in lakes (a) Oahe, (b) Sharpe,
(c) Francis Case, and (d) Lewis & Clark. Means within reservoirs with the
same letter are not significantly different (ANOVA with Tukey’s HSD test on
log10 transformed values; P < 0.05). Error bars represent SEs. Site codes for
Lake Oahe: BVB = Beaver Bay, BVC = Beaver Creek, CAR = Cannonball
River, CRC = Cheyenne River Confluence, CRU = Cheyenne River Upstream,
GRC = Grand River Confluence, GRU = Grand River Upstream, HAZ =
Hazelton, HTR = Heart River, KNR = Knife River, MIN = Minneconjou
Embayment, MOB = Mobridge Main Channel, MRC = Moreau River
Confluence, MRU = Moreau River Upstream, ODF = Oahe Dam Face, OKO =
Okobojo Embayment, SWC = Swan Creek Embayment, WPL = West Pollock
Embayment, WWE = West Whitlocks Embayment, and WWM = West
Whitlocks Main Channel. Site codes for Lake Sharpe: ANT = Antelope Creek,
BRC = Bad River Confluence, BRU = Bad River Upstream, FTG = Fort
George, HIP = Hipple Lake, JOE = Joe Creek, NSH = North Shore, SSB =
Sharpe Stilling Basin, and WEB = West Bend. Site codes for Lake Francis
Case: AMC = American Creek, CED = Cedar Shore Main Channel, NBE =
North Bay Embayment, NBM = North Bay Main Channel, PCE = Platte Creek
Embayment, PCM = Platte Creek Main Channel, WRC = White River
Confluence, and WRU = White River Upstream. Site codes for Lewis & Clark
Lake: GPD = Gavin’s Point Dam, LCE = Lewis & Clark Embayment, LCM =
Lewis & Clark Main Channel, NRC = Niobrara River Confluence, NRU =
Niobrara River Upstream, SPE = Springfield Embayment, and SPM =
Springfield Main Channel. ............................................................................... 40
xi Figure 2-4. Mean water Sr:Ca values at nine tributaries in Missouri River reservoirs
(Oahe, Sharpe, Francis Case, Lewis & Clark). Means with the same letter are
not significantly different (ANOVA with Tukey’s HSD test on log10
transformed values; P < 0.05). Error bars represent SEs. Site codes are as
follows: CAR = Cannonball River, CHR = Cheyenne River, GRR = Grand
River, HTR = Heart River, KNR = Knife R, MOR = Moreau River, BAR =
Bad River, WHR = White River, and NBR = Niobrara R. .............................. 41
Figure 2-5. Mean water Ba:Ca values at 20 collection sites in lakes (a) Oahe, (b) Sharpe,
(c) Francis Case, and (d) Lewis & Clark. Means within reservoirs with the
same letter are not significantly different (ANOVA with Tukey’s HSD test on
log10 transformed values; P < 0.05). Error bars represent SEs. Site codes for
Lake Oahe: BVB = Beaver Bay, BVC = Beaver Creek, CAR = Cannonball
River, CRC = Cheyenne River Confluence, CRU = Cheyenne River Upstream,
GRC = Grand River Confluence, GRU = Grand River Upstream, HAZ =
Hazelton, HTR = Heart River, KNR = Knife River, MIN = Minneconjou
Embayment, MOB = Mobridge Main Channel, MRC = Moreau River
Confluence, MRU = Moreau River Upstream, ODF = Oahe Dam Face, OKO =
Okobojo Embayment, SWC = Swan Creek Embayment, WPL = West Pollock
Embayment, WWE = West Whitlocks Embayment, and WWM = West
Whitlocks Main Channel. Site codes for Lake Sharpe: ANT = Antelope Creek,
BRC = Bad River Confluence, BRU = Bad River Upstream, FTG = Fort
George, HIP = Hipple Lake, JOE = Joe Creek, NSH = North Shore, SSB =
Sharpe Stilling Basin, and WEB = West Bend. Site codes for Lake Francis
Case: AMC = American Creek, CED = Cedar Shore Main Channel, NBE =
North Bay Embayment, NBM = North Bay Main Channel, PCE = Platte Creek
Embayment, PCM = Platte Creek Main Channel, WRC = White River
Confluence, and WRU = White River Upstream. Site codes for Lewis & Clark
Lake: GPD = Gavin’s Point Dam, LCE = Lewis & Clark Embayment, LCM =
Lewis & Clark Main Channel, NRC = Niobrara River Confluence, NRU =
Niobrara River Upstream, SPE = Springfield Embayment, and SPM =
Springfield Main Channel. ............................................................................... 42
Figure 2-6. Mean water Ba:Ca values at nine tributaries in Missouri River reservoirs
(Oahe, Sharpe, Francis Case, Lewis & Clark). Means with the same letter are
not significantly different (ANOVA with Tukey’s HSD test on log10
transformed values; P < 0.05). Error bars represent SEs. Site codes are as
follows: CAR = Cannonball River, CHR = Cheyenne River, GRR = Grand
River, HTR = Heart River, KNR = Knife R, MOR = Moreau River, BAR =
Bard River, WHR = White River, and NBR = Niobrara River. ...................... 43
Figure 3-1. Water and walleye sampling locations in Missouri River reservoirs. All sites
were sampled for water trace element concentrations using a syringe-filtration
protocol described in Shiller (2003). Shapes denote locations where fish of
xii different ages were sampled: right-angle triangles (age-0), upright triangles
(adult), diamonds (both)………………………………………………..……..74
Figure 3-2. Mean water (a) Sr:Ca and (b) Ba:Ca value at the 14 sites where both water
and age-0 Walleyes were collected in late summer 2012. Means with the same
letter are not significantly different (ANOVA with Tukey’s HSD test on log10
transformed values; P ≤ 0.05). Error bars represent 1
SEM…………………………………………………………………………76
Figure 3-3. Terminal otolith (a) Sr:Ca and (b) Ba:Ca ratios of age-0 Walleyes at the 14
sites where both water and age-0 Walleyes were collected in late summer
2012. Signatures represent mean terminal or adjusted mean terminal
concentrations that are temporally matched with water sampling. Means with
the same letter are not significantly different (ANOVA with Tukey’s HSD test
on log10 transformed values; P ≤ 0.05). Error bars represent 1 SEM .............. 77
Figure 3-4. Linear regression of age-0 Walleye terminal otolith (a) Sr:Ca and (b) Ba:Ca
signatures on water ratios at collection sites lakes Oahe, Sharpe, Francis Case,
and Lewis & Clark. Fish and water sampling occurred in late summer 2012.
Error bars represent 1 SEM. ............................................................................. 78
Figure 4-1. Water and walleye sampling locations in Missouri River reservoirs. All sites
were sampled for water trace element concentrations using a syringe-filtration
protocol described in Shiller (2003). Shapes denote locations where fish of
different ages were sampled: right-angle triangles (age-0), upright triangles
(adult), diamonds (both) ……………………….....…………………………116
Figure 4-2. A conceptual model for evaluating movement and entrainment of walleye in
Missouri River reservoirs. Site assignment of adult walleye to locations in the
Missouri River resembled a decision tree involving synthesis of bivariate
water and otolith signatures, the river-wide water-otolith relationship,
collection locations, and reservoir orientation. The water-otolith relationship
generated from mean terminal otolith signatures of age-0 walleye collected in
fall 2012. ………………………..........................................................……117
Figure 4-3. Mean water (a) Sr:Ca and (b) Ba:Ca value at the 14 sites where both water
and age-0 Walleyes were collected in late summer 2012. Means with the same
letter are not significantly different (ANOVA with Tukey’s HSD test on log10
transformed values; P ≤ 0.05). Error bars represent 1
SEM.………………………...................................................................……118
xiii Figure 4-4. Linear regression of age-0 walleye mean terminal otolith (A) Sr:Ca and (B)
Ba:Ca signatures on water ratios at collection sites lakes Oahe, Sharpe,
Francis Case, and Lewis & Clark, North Dakota and South Dakota, USA. Fish
and water sampling occurred in fall 2012. Error bars represent 1
SEM.………………………...................................................................……119
Figure 4-5. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye
collected in Lake Sharpe, South Dakota, USA in Summer 2013. Site
assignments reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some
adults (e.g., black circle) displayed relatively stable element:Ca ratios
indicating prolonged site residence, whereas others (e.g., white triangle)
exhibited variable signatures suggesting intra-reservoir movement and
entrainment through mainstem dams. All transects proceed from the core of
the otolith toward the edge. Individuals depicted here were collected at Sharpe
Stilling Basin. ……………………............................................................……120
Figure 4-6. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye
collected in Lake Francis Case, South Dakota, USA in Summer 2013. Site
assignments reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some
adults (e.g., black triangle) displayed relatively stable element:Ca ratios
indicating prolonged site residence, whereas others (e.g., white triangle)
exhibited variable signatures suggesting intra-reservoir movement and
entrainment through mainstem dams. All transects proceed from the core of
the otolith toward the edge. Individuals depicted here were collected near
Chamberlain, South
Dakota.………………………....................................................................……121
Figure 4-7. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye in
Lewis & Clark Lake, South Dakota, USA in Summer 2013. Site assignments
reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., white
circle) displayed relatively stable element:Ca ratios indicating prolonged site
residence, whereas others (e.g., black circle) exhibited variable signatures
suggesting intra-reservoir movement and entrainment through mainstem dams.
All transects proceed from the core of the otolith toward the edge. Individuals
depicted here were collected at Gavin’s Point
Dam.………………………........................................................................……122
Figure 5-1. Missouri River watershed and location of the Lewis and Clark Delta along the
South Dakota-Nebraska border. .....................................................................148
xiv Figure 5-2. Gauge height in the Lewis and Clark Delta, South Dakota from 1980-2012.
Flooding in 2011 represented the most severe hydrological disturbance in
recent history. Gauge height was measured at a United States Geological
Survey site in Springfield, SD (USGS 06466700, Latitude 42°51'21",
Longitude 97°53'06"). ....................................................................................149
Figure 5-3. Nonnetric multidimensional scaling (NMS) ordinations of catch-per-uniteffort (CPUE) for juvenile native and sport fish sampled during pre-flood
(2004-2010), flood (2011), and post-flood (2012) periods (symbols) in the
Lewis and Clark Delta, South Dakota. Multivariate heterogeneity is explained
by directional changes in CPUE of fishes categorized by introduction history
(Introduced, Native, Sport), trophic guild (D = detritivore; O = omnivore; P =
piscivore), and species (CAP = common carp; FRD = freshwater drum; RIC =
river carpsucker; BGL = bluegill; BLC = black crappie; SMB = smallmouth
bass; WHC = white crappie). NMS was performed in Primer (PRIMER-E Ltd,
Plymouth, United Kingdom; Clarke and Gorley 2006) using Bray-Curtis
ecological distance. Sample sizes are included in Table 5-1. ........................150
Figure 5-4. Nonmetric multidimensional scaling (NMS) ordinations of catch-per-uniteffort (CPUE) for adult native and sport fish and small-bodied native species
sampled during pre-flood (2004-2010), flood (2011), and post-flood (2012)
periods (symbols) in the Lewis and Clark Delta, South Dakota. Multivariate
heterogeneity is explained by directional changes in CPUE of fishes
categorized by introduction history (Introduced, Native, Sport, Small-bodied
Native [SBN]), trophic guild (D = detritivore; I = Introduced; O = omnivore; P
= piscivore), and species (CAP = common carp; FRD = freshwater drum; RIC
= river carpsucker; SHR = shorthead redhorse; BGL = bluegill; CCF =
channel catfish; LMB = largemouth bass; SAR = sauger; EMS = emerald
shiner; RES = red shiner; SFS = spotfin shiner). NMS was performed in
Primer (PRIMER-E Ltd, Plymouth, United Kingdom; Clarke and Gorley
2006) using Bray-Curtis ecological distance. Sample sizes are included in
Table 5-1. .......................................................................................................151
Figure 5-5. Pre-flood (June 2010) and post-flood (October 2011) frequency of channels,
backwaters, and sandbars in the Lewis & Clark Delta by equidistant transect
between the Niobrara River confluence and downstream limit of the
Delta... ............................................................................................................152
xv LIST OF TABLES
Table 2-1. Morphometric and hydrological features of North Dakota and South Dakota
Missouri River reservoirs. Modified from Erickson et al. (2008)................... 36
Table 3-1. Results of one-way ANOVA testing for spatial variability in water element:Ca
ratios among Missouri River tributaries and sites within reservoirs. A
significant P value (≤ 0.05) indicates water chemistry is spatially variable in
the associated water body. ...............................................................................72
Table 3-2. Results of k-sample nearest neighbor discriminant analysis with leave-one-out
jackknife cross-validation using Sr:Ca and Ba:Ca ratios of age-0 Walleye. Data
include the number known to have come from site types and individual sites
(Known), the number assigned to those locations (Assigned), the percent
deviation of Assigned from Known (Deviation), and the percentage of
individuals classified to known locations (Accuracy). Signatures were
measured at otolith locations synchronized with water sample collection to
ensure reliable spatial blueprinting.. ................................................................ 73
Table 3-3. Percent (number) natal contribution of tributaries and mainstem/embayment
locations to adult Walleye populations in Missouri River reservoirs, North
Dakota and South Dakota, USA, grouped by collection reservoir and site. FC
and LC denote Lake Francis Case and Lewis & Clark Lake, respectively. ..... 74
Table 4-1. Results of one-way ANOVA testing for spatial variability in water element:Ca
ratios among Missouri River tributaries and sites within reservoirs. A
significant P value (≤ 0.05) indicates water chemistry is spatially variable in
the associated water body.……………………………….………………….110
Table 4-2. Results of k-sample nearest neighbor discriminant analysis with leave-one-out
jackknife cross-validation using Sr:Ca and Ba:Ca ratios of age-0 Walleye. Data
include the number known to have come from site types and individual sites
(Known), the number assigned to those locations (Assigned), the percent
deviation of Assigned from Known (Deviation), and the percentage of
individuals classified to known locations (Accuracy). Signatures were
measured at otolith locations synchronized with water sample collection to
ensure reliable spatial blueprinting.. .............................................................. 111
Table 4-3. Percent (number) of adult walleye exhibiting downstream movement,
upstream movement, and site residency at reservoir (Oahe, Sharpe, Francis
Case, Lewis & Clark) and statewide (North Dakota [ND], South Dakota [SD])
spatial scales. At the statewide scale, “downstream” and “upstream” denote
movements to SD and ND, respectively.….. .. .............................................. 112
xvi Table 4-4. Summary of adult walleye entrainment in Missouri River reservoirs, where %
Flood-entrained represents the percentage of all fish and entrained fish (in
parentheses) that were entrained during the 2011 flood. Similarly, % Baseline
represents the percentage of all fish and entrained fish that were not entrained
during the 2011 flood... .................................................................................. 111
Table 4-5. Summary of adult walleye entrainment by collection site in Missouri River
reservoirs. Entrained (flood) represents all entrained and 2011 flood-entrained
individuals (in parentheses). GPD denotes Gavin’s Point Dam. ................... 113
Table 5-1. Fish species collected from 2004 – 2012 in the Lewis and Clark Delta
organized by introduction history (SBN = small-bodied native) and trophic
guild. Sample sizes of fishes captured by electrofishing (EF) are listed by
length category (adult, juvenile/small-bodied). Only juvenile fishes were
sampled with mini-fyke netting (MF)……………………………………….145
Table 5-2. Sampling distribution of fishes collected during flood periods (pre-flood
[2004-2010], flood [2011], post-flood [2012]) in the Lewis and Clark Delta.
Fishes are organized by introduction history, trophic guild, and gear type.
Electrofishing (EF) data included adults in all categories and guilds except
small-bodied native (SBN), invertivore, and planktivore, which represent
juvenile and small-bodied individuals. EF was not conducted during the flood.
Mini-fyke net (MF) data included juvenile individuals in all categories and
guilds except those mentioned above............................................................. 146
Table 5-3. Structural indices (species richness, evenness [J’], diversity [Fisher’s α]) of
fishes collected during flood periods (pre-flood [2005, 2008-2009], post-flood
[2012]) in the Lewis and Clark Delta. The entire species pool was reduced to
common species (> 1% of total catch during a particular sampling event) to
remove effects of rare species and species difficult to collect. Letters indicate
significant differences among flood periods. ................................................. 147
xvii ABSTRACT
RAPID RESPONSE TO A CATASTROPHIC FLOOD: EFFECTS ON AQUATIC
RESOURCES IN MISSOURI RIVER RESERVOIRS
ANDREW KENNETH CARLSON
2015
Ecological effects of an historic flood in the Missouri River in 2011 are largely
unknown. In rapid response, we assessed walleye Sander vitreus environmental history in
Missouri River reservoirs using otolith microchemistry and evaluated flood effects on the
Lewis & Clark Delta fish community and aquatic habitats.
Geologic heterogeneity drove spatial variability in water chemistry throughout
impoundments. Otolith trace element concentrations of age-0 and adult walleyes varied
spatially in accordance with water chemistry. Bivariate natal signatures of adults
exhibited site-specific heterogeneity sufficient to accurately identify natal tributaries and
mainstem/embayment sites. The Moreau and Cannonball rivers were particularly
important natal sites in Lake Oahe, as were North Shore, West Bend, Platte Creek, and
the Lewis & Clark Delta in southern reservoirs.
Walleye movement within and among Missouri River reservoirs was dominated
by downstream passage and site residency before, during, and after the flood; upstream
movement was relatively uncommon. Entrainment was most extensive during the flood
and increased progressively moving downstream. Otolith microchemistry indicates
divergent entrainment among hydrologic regimes, connectivity of walleye populations
among impoundments, and the importance of collaborative riverscape management
across state boundaries for walleye management in Missouri River reservoirs.
xviii Fish community evenness (J’) in the Lewis and Clark Delta decreased from 0.882
to 0.725 before the flood but rose to 0.835 after the disturbance. Pre-flood species
richness declined from 25 to 15, whereas post-flood richness stabilized above 20.
Diversity (Fisher’s α) decreased from 4.56 to 3.48 before the flood and stabilized at 3.27
after the disturbance. A majority of species were more abundant after the flood, whereas
juvenile freshwater drum Aplodinotus grunniens and white crappie Pomoxis annularis
were less abundant and small-bodied native species remained stable. Channel and
backwater frequency and main channel width in the Delta declined after the flood,
whereas sandbars became more abundant. Habitat changes likely had minimal effects on
the fish community due to persistence of shallow-water habitats important for spawning
and rearing. Our results suggest the Delta serves as a refuge environment for juvenile and
adult fishes during and immediately after catastrophic flooding.
1 CHAPTER 1. INTRODUCTION
Large floodplain rivers are among the world’s most complex ecosystems
(Dynesius and Nilsson 1994). Meandering sequences of riffles, runs, and pools change
rapidly and disappear at localized spatial scales but persist at broader dimensions of space
and time, creating a dynamic equilibrium (Galat et al. 1996) that generates local volatility
and global stability in biological community structure. The Flood Pulse Concept (Junk et
al. 1989) summarizes complex structural and functional properties of large floodplain
rivers. It theorizes aquatic biomass in unaltered systems is derived from floodplain
production rather than downstream transport of organic matter. Thus, lateral riverfloodplain connectivity and nutrient exchange exert more control over biotic processes
than longitudinal upstream-downstream linkages. Seasonal floods provide lateral
connectivity, inundating floodplains with nutrient-rich water (Stanford et al. 1996) and
controlling population and community dynamics of aquatic organisms (Kingsford 2000).
Despite their ecological importance, floods cause social and economic costs that have
driven river management and flood control throughout human history.
River Management
For centuries humans have altered rivers for societal benefits, including water
supply, navigation, and flood control. These goods and services promote human wellbeing and economic growth. In ecological terms, river management modifies natural flow
regimes and trophic relationships, causing ecosystem changes. Fluvial habitat-forming
processes are diminished or eliminated as connectivity among channel, floodplain, and
upland environments is reduced. Existing habitats are homogenized by interactive forces
2 (e.g., temperature shifts, channel simplification, floodplain desiccation, etc.) that remove
native species and enable non-native organisms to flourish (Stanford et al. 1996).
Flow regulation in the form of dams, wing dykes, levees, and other structures is
the most pervasive anthropogenic alteration to rivers worldwide (Standford et al. 1996).
Globally, 172 of 292 large river systems (i.e., virgin mean annual discharge, VMAD,
greater than 350 m3s-1) are fragmented by dams (Nilsson et al. 2005), which increased
sevenfold between 1950 and 1986 (Dynesius and Nilsson 1994). In the northern third of
the world, all 139 large river systems are regulated by humans (Dynesius and Nilsson
1994; Kingsford 2000). More than 45,000 large dams (> 15 m high; Nilsson et al. 2005)
and 800,000 small dams (Johnson 2002) fragment rivers throughout the world, with
79,000 constructed in the United States (ASCE 2005). Flow regulation diminishes
biological production and biodiversity in riverine ecosystems by altering dynamic
interactive processes of the river continuum (Stanford et al. 1996). Ecological changes
resulting from flow regulation have emerged throughout the world.
International research provides a global perspective for conceptualizing Missouri
River management. Australian rivers (Gehrke et al. 1995) and floodplain wetlands
(Kingsford 2000) provide valuable ecosystem goods and services and support abundant,
diverse populations of birds, fishes, invertebrates, aquatic plants, and microbes. Dams
and weirs in these ecosystems depress flood frequency, fragment and homogenize
habitats, cause population declines of native species, and depress biodiversity by severing
river-floodplain connectivity. Unnatural flow regimes desynchronize reproductive
rhythms of native species from environmental cycles, effectively opening niches for
generalist species such as common carp Cyprinus carpio. River management produced
3 analogous effects in the Upper Paraná River in southern Brazil, where hydrological
alterations disrupted evolutionary equilibria among natural flow regimes and fish life
history processes (e.g., gonad maturation, foraging, spawning, and larval development;
Agostinho et al. 2004).
River management continues to alter macroinvertebrate (Robinson et al. 2011),
fish (Mims and Olden 2013), and bird (Jonsson et al. 2012) populations throughout the
world. Fish communities often homogenize and native populations decline as
impoundment and channelization fragment habitat and hinder or prevent movement. Yet
reservoirs support multi-million dollar sport fisheries and recreation industries that
provide substantial societal benefits. As such, effective reservoir management is
important for biota and humans alike. The Missouri River provides an informative case
study for examining opportunities and challenges for aquatic resource management in
impounded river systems.
Opportunities and Challenges for Missouri River Management
Regulated rivers provide substantial societal benefits, including water supply,
flood control, power generation, transport of goods, irrigation for agriculture, and
recreational opportunities (Allen et al. 2008). These goods and services generate
economic growth and meet diverse commercial and consumer demands (Niemi and
Raterman 2008). Competition for reservoir goods and services adds layers of complexity
to a reservoir management paradigm that was formerly dominated by biological concerns
(Miranda 1996), which are intricate and controversial themselves. As the human
dimensions of reservoirs become more prominent and imperative, management must
4 evolve to keep pace with both biological and social demands.
Following the Lewis and Clark expedition from 1804 to 1806, the Missouri River
became a path for European settlement and trade as the nation expanded westward.
Catastrophic flooding was relatively common in the late 1800s and early 1900s,
generating significant societal and economic costs (Erickson et al. 2008). Combined with
the nation’s growing interest in using the Missouri River for navigation and hydroelectric
power in the 1930’s and 1940’s, flooding prompted river management activities that
continue today. Impoundment of the Missouri River began in 1933 with the construction
of Fort Peck Dam and Fort Peck Reservoir (Erickson et al. 2008). In 1944, the U.S. Army
Corps of Engineers Flood Control Act commissioned the creation of five more reservoirs
(i.e., Sakakawea, Oahe, Sharpe, Francis Case, and Lewis and Clark) that were completed
between 1952 and 1963. Missouri River reservoirs have since provided substantial goods
and services, including flood control, irrigation, navigation, hydropower, fish and wildlife
resources, and recreation (Erickson et al. 2008).
Despite providing substantial societal contributions, Missouri River reservoirs
present challenges for aquatic resource management. In 2000, passage of comprehensive
federal environmental legislation initially sanctioned in the 1960s and 1970s (Endangered
Species Act 2000; Federal Water Pollution Control Act 2000; National Environmental
Policy Act 2000; Wild and Scenic Rivers Act 2000) added complexity to an already
complicated, controversial situation (Erickson et al. 2008). Moreover, disturbances in
Missouri River reservoirs are unpredictable and difficult to manage. For instance, largescale flooding causes entrainment, an occurrence in which aquatic organisms are flushed
downstream through dams, weirs, and other structures. Entrainment has crucial
5 implications for fisheries management in Missouri River reservoirs (e.g., source-sink
dynamics), but managers have little information regarding the magnitude and extent of
fish passage, impairing their ability to anticipate and mitigate consequences of future
disturbances. Reliable methods to quantify entrainment are needed to sustain ecologically
and economically important fish populations in Missouri River reservoirs.
Sedimentation and aquatic invasive species also present challenges for Missouri
River management. Sediment accumulation fragments aquatic habitats (Walburg 1976,
1977; Patton and Lyday 2008) and reduces storage capacity of reservoirs (Coker 2000).
However, freshwater deltas created by sediment deposition provide riverine habitat that
supports higher fish species diversity than lotic areas of reservoirs (Kaemingk et al.
2007). In the future, Missouri River managers will be faced with striking a balance
between costs and benefits of sedimentation. Aquatic invasive species such as zebra
mussel Dreissena polymorpha veligers have been found in Lake Francis Case and the
riverine sections of the Missouri River downstream of Lake Francis Case and Lewis and
Clark (Erickson et al. 2008). In addition, bighead carp Hypophthalmichthys nobilis and
silver carp H. molitrix have been collected from the Missouri River below Lewis and
Clark Lake. There are concerns that these species will exert deleterious effects on native
riverine fishes and sport species if they became established in mainstem reservoirs.
Finally, Missouri River managers will face challenges implementing an adaptive
management paradigm that is sufficiently flexible to respond to political, social, and legal
changes in the reservoir management environment (Erickson et al. 2008).
Missouri River impoundments furnish societal benefits as well as challenges for
aquatic resource management. In the midst of a complex management environment,
6 scientific inquiry and multiagency collaboration will continue to illuminate strategies for
preserving and enhancing multiple reservoir uses. The promise of success makes
reservoir management an intriguing area of research. With this vision in mind, we review
specific opportunities and challenges for aquatic resource management in Missouri River
reservoirs in response to a catastrophic flood in 2011.
Catastrophic Flooding
In 2011, late-melting snow and late-spring rainfall in the Rocky Mountains and
throughout the Great Plains caused catastrophic flooding in the Missouri River. Runoff
totaled 75.2 billion m3 and exceeded system capacity by > 20 % above Sioux City, IA
(USACE 2012). All six mainstem reservoir dams released record amounts of water.
Citizen evacuations occurred in many cities as floodwaters damaged residential,
corporate, industrial, agricultural, and transportation infrastructure (Grigg et al. 2011).
The Federal Emergency Management Agency provided more than $395 million in
disaster relief (Freeman and Jones 2012), and the USACE incurred $1 billion in damage
expenses. Direct and indirect costs to municipalities, families, and individuals are
unknown and likely far greater (Grigg et al. 2011).
In addition to social, economic, and hydrological effects of the flood, the
disturbance caused ecological changes. Preliminary research indicates substantial
physical alterations, including bed degradation, thalweg migration, and loss of riverfloodplain connectivity (Cowman 2012). Further geomorphological research is essential
to fill knowledge gaps and lay a foundation for assessing biotic effects. Qualitative
evidence suggests extensive fish passage through hydroelectric facilities statewide and
7 modifications to aquatic habitats in freshwater deltas. With uncertainties regarding
ecological consequences of the flood, these topics represent promising research avenues.
Entrainment
Entrainment is a phenomenon whereby aquatic organisms are washed
downstream through dams, weirs, and other structures and discharged back into the
environment (Nowakowski et al. 2006). It has been implicated as a major source of fish
mortality (OTA 1995; Cada et al. 1997) and may hinder efforts to maintain reproductive
populations and recreational fisheries. Although it occurs across a range of discharge
levels during flood and non-flood periods, entrainment is most extensive when discharges
are high (Walburg 1971; Unkenholz 1998; GeoSense 2011). Entrainment has been
extensively researched in the western United States as it causes population declines of
highly prized, economically important salmonid species (Clothier 1953; Hallock and Van
Woert 1959; Coutant and Whitney 2000; Schrank and Rahel 2004; Carlson and Rahel
2007; Gale et al. 2008; Roberts and Rahel 2008; Prince 2009; GeoSense 2011; Simpson
and Ostrand 2012). However, entrainment research in the central United States is scarce,
especially for resident reservoir species (Walburg 1971; Navarro and McCauley 1996;
Unkenholz 1998).
A number of interactive factors influence entrainment of fishes in reservoirs.
Reservoir size (GeoSense 2011), elevation (Unkenholz 1998), volume (GeoSense 2011),
discharge (Shreffler et al. 1994; Coutant and Whitney 2000; Ploskey et al. 2005), dam
intake and thermocline depth (Unkenholz 1998; Nowakowski et al. 2006), and season and
time of day (Burczynski et al. 1987; Unkenholz 1998; Nelson-Stastny 2001; Jaeger et al.
8 2005; Spinelli 2010) affect water movement, fish behavior, and entrainment.
Characteristics such as species (Nowakowski et al. 2006), life stage (Unkenholz 1998),
year class strength (Grimaldo et al. 2009), and vertical distribution (Burczynski et al.
1987; Unkenholz 1998; Nelson-Stastny 2001; Jaeger et al. 2005; Spinelli 2010) also
influence entrainment susceptibility. Walleye Sander vitreus and sauger Sander
canadensis exhibit elevated entrainment rates during high discharges (Walburg 1971;
Jaeger et al. 2005; GeoSense 2011), which is problematic as they are prized recreational
species throughout the Midwest. Entrainment causes injury and mortality, prevents access
to essential habitats, precludes genetic exchange among populations, increases
hybridization, and impairs recruitment (Larinier 2000; Nowakowski et al. 2006).
Although entrainment may be mitigated using physical (e.g., fish screens, angled bar
racks, louvers, and barrier nets) and behavioral (e.g., light, sound, air bubble currents, and
electricity) barriers, these methods are impractical on large scales and are limited to areas
where entrainment is known to occur.
Entrainment is difficult to study due to diverse causative factors and biased
estimation methods. Traditional techniques such as hydroacoustics, telemetry, and netting
(Unkenholz 1998) are prone to limitations such as cost and impracticality that prevent
reliable estimation. In recent years, researchers have used microchemical methods to
investigate environmental history and entrainment of marine and freshwater fishes.
Otoliths, dentary bones, fin rays, and other calcified structures accumulate permanent
trace element depositions in proportion to water column concentrations (Campana and
Thorrold 2001; Zeigler and Whitledge 2011; Wolff et al. 2012) and accrete aragonitic
calcium in daily concentric rings that construct a permanent growth record (Campana and
9 Thorrold 2001). Association of calcified structure biochronology with elemental
composition permits retrospective description of environmental history (e.g., natal
origins, movement, entrainment) when fishes have resided in chemically distinct
locations for a period of time sufficient to incorporate unique elemental signatures
(Kennedy et al. 2002).
Otolith microchemistry has facilitated diverse environmental history research on
fishes. Scientists have examined natal origins (Zeigler and Whitledge 2010, 2011; Wolff
et al. 2012), life history movements (Brenkman et al. 2007; Allen et al. 2009), stock
composition (Bickford and Hannigan 2005; Coghlan et al. 2007), habitat fingerprinting
(Brazner et al. 2004), and invasion biology of nonnative species (Munro et al. 2005;
Whitledge 2009). However, otolith microchemistry is rarely employed for entrainment
research. During a pelagic organism decline in the San Francisco Bay Delta, otolith
signatures provided a “geographic scope” for conservation of federally threatened delta
smelt Hypomesus transpacificus, yielding critical information on entrainment, growth,
and recruitment (Hobbs et al. 2007; Hobbs 2010). Otolith microchemistry has also been
used to evaluate entrainment of bull trout Salvelinus confluentus (Prince 2009) and
common triplefins Forsterygion lapillum (Shima and Swearer 2009). With considerable
knowledge gaps regarding entrainment in the central United States, otolith
microchemistry is a promising tool across many environments and species.
Sedimentation
Many large-river impoundments in the United States and throughout the world
were constructed in the 1950s and 1960s (Avakyan and Lakovleva 1998; Miranda et al.
2010). Although reservoirs were designed to accumulate sediment slowly (Palmieri et al.
10 2001), symptoms of aging (e.g., sedimentation, shoreline erosion) have emerged.
Sedimentation fragments aquatic habitats (Walburg 1976, 1977; Patton and Lyday 2008),
reduces storage capacity (Coker 2000) and waterfront access (Elliott and Jacobson 2006;
USACE 2006), causes localized flooding (USACE 2001), depresses productivity (Wood
2010), and decreases fish production (Slipke et al. 2005). Sediments also form freshwater
deltas at mouths of tributaries and upper reaches of reservoirs (Johnson 2002). Although
deltas may exacerbate consequences of reservoir aging, they contain warm, turbid water
and heterogeneous habitats that simulate historic riverscapes (Graeb et al. 2009) and
support diverse fish communities (Kaemingk et al. 2007). The six mainstem Missouri
River reservoirs have expanding freshwater deltas with associated costs and benefits. The
USACE is examining feasibility of sediment management alternatives (e.g., dewatering
and sediment flushing, channelization, dredging, tributary diversion, downstream
sediment diversion) to mitigate reservoir aging (USACE 2011a, b). As it renders
decisions in the future, it will be imperative for the agency to balance ecological services
of deltas with benefits of sediment management.
Project Overview
Regulated rivers provide societal benefits and present challenges for aquatic
resource management. Floods, entrainment, and sedimentation pose potential hindrances
for management, yet scientific inquiry and multiagency collaboration continue to
facilitate conservation successes. The 2011 Missouri River flood provided a rare
opportunity to unite distinct research questions concerning aquatic resources in North
Dakota and South Dakota. Throughout South Dakota and the lower half of North Dakota,
11 we evaluated otolith microchemistry as a tool for examining environmental history (e.g.,
natal origins, movement, entrainment) of walleye in Missouri River reservoirs. A
requirement of this technique is spatial variability in water chemistry that permits
retrospective assignment of individuals to specific sites. An extensive network of
tributaries flow into the Missouri River through areas with distinct geology. North
Dakota and South Dakota are characterized by spatial variability in surficial geology
caused by Pleistocene glaciation east of the Missouri River 10,000–12,000 years ago.
Geologically young glacial sediments predominate east of the river, whereas older
Cretaceous sandstones, clays, and shales exhibit a north-south gradient in west of the
river. Geologic heterogeneity drives spatial variability in water chemistry and renders
otolith microchemistry a viable technique for evaluating fish environmental history in
Missouri River reservoirs.
A principal research objective of this project was to examine intra- and interreserovir movement of walleye. The South Dakota Department of Game, Fish and Parks
(SDGFP) observed extensive entrainment through Missouri River dams during the flood,
but the magnitude and extent of fish passage were not quantified prior to this study.
Although certain walleye spawning and recruitment areas were identified previously,
knowledge gaps (e.g., relative tributary contribution) remained to be filled. We addressed
these and related environmental history questions using otolith microchemistry. This
research fostered development of a predictive tool for quantifying entrainment across a
range of discharge levels during flood and non-flood periods, providing agencies with
valuable information for science-based management.
12 On a localized scale, we investigated effects of flooding on aquatic resources in
the Lewis and Clark Delta. Formed by sediments from the Niobrara River, this unique
ecosystem simulates historic Missouri River habitat and supports higher richness and
diversity of fishes than lentic portions of the reservoir (Kaemingk et al. 2007). Treating
the 2011 flood as a natural experiment, we compared fish community data and georeferenced aquatic habitat information from pre-flood and post-flood periods to assess
ecological effects of flooding in the Delta. This study revealed critical information about
disturbance ecology in the Missouri River that may assist the USACE and SDGFP in
aquatic resource management. Coupled with otolith microchemistry information, this
research will illuminate strategies for addressing challenges in aquatic resource
management in Missouri River reservoirs.
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25 CHAPTER 2: SPATIAL HETEROGENEITY IN TRACE ELEMENT CHEMISTRY IN
MISSOURI RIVER RESERVOIRS
This chapter is being prepared for submission to Lake and Reservoir Management and
was co-authored by Mark J. Fincel, Christopher M. Longhenry, and Brian D. S. Graeb.
26 Abstract
Concentrations of chemical elements in freshwater bodies reflect surficial and
bedrock geology and weathering in surrounding watersheds. Missouri River reservoirs
drain areas with heterogeneous geology, but water column chemistry has not been
comprehensively examined. We evaluated spatial variability in trace element
concentrations in tributaries and mainstem/embayment sites in four Missouri River
reservoirs (Oahe, Sharpe, Francis Case, Lewis & Clark) in North Dakota and South
Dakota, USA. Duplicate water samples from 57 locations were collected in July 2012 and
February 2014 and analyzed using high-resolution inductively coupled plasma mass
spectrometry. Spatial and temporal patterns in water chemistry were assessed using
analysis of variance on log10 transformed concentrations. Sr:Ca, Ba:Ca, Mg:Ca, Mn:Ca,
and Na:Ca were spatially variable, and Sr:Ca and Ba:Ca temporally consistent, among
reservoirs and tributaries. Tributaries exhibited greater heterogeneity in element:Ca ratios
than embayment and mainstem sites. Spatial heterogeneity reflected variability in
geology and wreathing among tributaries, embayments, and mainstem environments.
This research enhances understanding of geological influences on water chemistry in
Missouri River reservoirs and facilitates future studies that necessitate spatially variable
trace element blueprints, including calcified structure microchemistry of fishes.
Key words: strontium, barium, calcium, Missouri River, North Dakota, South Dakota
27 Introduction
Concentrations of chemical elements in water bodies throughout the world reflect
three major mechanisms: atmospheric precipitation, rock weathering, and evaporation
(Gibbs 1970). The relative importance of these processes varies with salinity and
precipitation/runoff, with freshwater bodies in the United States dominated by rock
weathering. Trace elements (e.g., strontium [Sr[, barium [Ba], magnesium [Mg],
manganese [Mn], sodium [Na]) are defined as stable and/or radioactive elements with
concentrations < 1 mg/L (i.e.., 10-5 M solution for elements having a relative atomic
weight of 100; Beneš 1980). Rivers of the United States that drain watersheds with
distinct geology exhibit spatial variability in trace element chemistry. Quantifying spatial
and temporal patterns in trace element chemistry permits understanding of geological
influences on riverine water chemistry and lays a foundation for additional studies that
necessitate trace element blueprints. For example, contemporary fish environmental
history research links trace element concentrations in calcified structures (e.g., otoliths
[Gahagan et al. 2012], fin rays [Phelps et al. 2012]) with those in the water column to
retrospectively examine natal origins and movement.
Trace element chemistry in the Missouri River, USA, is influenced by the distinct
geological characteristics of tributaries, embayments, and mainstem environments.
However, knowledge of spatial and temporal patterns in trace element is incomplete. In a
previous study (Phelps et al. 2012), water samples were collected in the Lower Missouri
River (rkm 45.4) and the Middle Mississippi River during spring, summer, and fall 2006–
2008. Water samples were also obtained from Lake Oahe during 2008, and combined
data indicated that water strontium:calcium (Sr:Ca) ratios were higher in upstream
28 portions of the Missouri River compared with both the lower Missouri River and the
Mississippi River (Zeigler 2009). However, relative differences in trace element
signatures within and among Lakes Oahe, Sharpe, Francis Case, Lewis & Clark have not
been measured. This degree of spatial resolution is necessary for understanding smallscale geological influences on water chemistry and addressing broader questions (e.g.,
fish environmental history) germane for reservoir management. Our objective was to
evaluate spatial and temporal patterns in trace element chemistry within and among North
Dakota and South Dakota Missouri River reservoirs, particularly tributaries. We
hypothesized that element:Ca signatures would vary across space but not time in
accordance with spatial heterogeneity and temporal consistency in geology.
Study Site
The Missouri River is the longest river in North America, flowing 3,768 km from
Brower’s Spring, Montana to its confluence with the Mississippi River at St. Louis,
Missouri. It spans 1,371,017 km2, the second-largest riverine drainage area in the United
States, and encompasses 47 tributaries with drainage basins greater than 1000 km2 (Galat
et al. 2005). Much of the Missouri River lies within the semi-arid Great Plains, with half
of the basin receiving less than 41 cm/yr of precipitation (Galat et al. 2005). Average
July–October water temperatures range from 21.5 C to 27.0°C in most of the river, but
temperatures below reservoirs are depressed due to hypolimnetic water releases (Galat et
al. 2001). Cropland (37%) and grassland (30%) dominate land use within the basin,
which is minimally developed by humans (9%; Revenga et al. 1998). The river in North
Dakota and South Dakota is impounded into four reservoirs (Table 2-1) with
29 heterogeneous geology (Fig. 2-1). Cenozoic glacial sediments (e.g., Illinois and
Wisconsin glacial sediments) predominate east of the Missouri River, whereas older
Mesozoic sediments (e.g., sandstonces, shales, clays) exhibit a north–south gradient west
of the river.
Materials and Methods
Duplicate water samples from 57 locations in North Dakota and South Dakota
Missouri River reservoirs were collected in summer 2012 and winter 2014 to assess
spatial and temporal patterns in trace element concentrations. A reliable syringe filtration
method for sampling in remote field locations (Shiller 2003) was employed. All water
samples were collected with nitrile gloves and stored in sealed coolers (i.e., no light
penetration) before filtration. Sampling sites included main channels, embayments,
stilling basins, tailraces, and free-flowing tributaries. Trace element (i.e., 88Sr, 137Ba,
24
Mg, 55Mn, and 23Na) and calcium (43Ca) concentrations of water samples were
quantified using high-resolution inductively coupled plasma mass spectrometry (HRICPMS) at the University of Southern Mississippi. Spatial patterns in water chemistry in
summer 2012 and temporal patterns between summer 2012 and winter 2014 were
assessed at the reservoir and tributary scales using analysis of variance (ANOVA) with
Tukey’s Honestly Significant Difference (HSD) test for multiple comparisons on log10transformed water data to ensure normality and homoscedasticity.
Results
30 Tributaries on the western side of the Missouri River exhibited a decreasing
north-south Sr:Ca gradient (Fig. 2-2) consistent with latitudinal variation in surficial
geology. Mainstem sites on the eastern side of the river generally had more consistent
Sr:Ca ratios. Ba:Ca ratios of western tributaries followed the opposite north-south
pattern, increasing moving downstream, but eastern mainstem sites were relatively
homogenous. All element:Ca ratios were spatially heterogeneous at reservoir and
tributary scales (Table 2-2) in the same way each year for Sr:Ca (F1,179 = 69.89, P = 0.10;
Fig. 2-3, 2-4) and Ba:Ca (F1,179 = 102.62, P = 0.49; Fig. 2-5, 2-6) and different ways
annually for Mg:Ca (F1,179 = 20.69, P < 0.01), Mn:Ca (F8,6 = 3614.60, P < 0.01), and
Na:Ca (F1,103 = 45.50, P < 0.01).
Discussion
The chemistry of water bodies throughout the world is regulated by diverse
processes (i.e., atmospheric precipitation, rock weathering, evaporation), with rock
weathering predominating in freshwater bodies (Gibbs 1970). As such, spatial variability
in surficial and bedrock geology and weathering rates causes chemical differences within
and among United States rivers, as documented herein for the Missouri River in North
Dakota and South Dakota. Rock weathering and soil formation are governed by diverse
factors (e.g., trace element solubility, water pH, adsorption characteristics, hydration,
coprecipiation, colloidal dispersion, complexation; Durum and Haffty 1961) that
collectively caused spatial differences in element:Ca ratios among tributaries,
embayments, and mainstem environments in the present study. Anthropogenic (e.g,.
mining, manufacturing) and biotic (e.g., vegetation) factors may also have influenced
31 water chemistry in Missouri River reservoirs, but to a lesser degree than weathering.
Sr:Ca and Ba:Ca concentrations were temporally consistent as in prior studies (Gahagan
et al. 2012, Phelps et al. 2012), whereas Mg:Ca, Mn:Ca, and Na:Ca ratios were unstable
over time, confirming previous research (Wetzel 1983) and likely reflecting complex
biogeochemistry (Marshall 1979) in freshwater environments. These results are
informative from both scientific and applied management perspectives. Missouri River
water chemistry has not been previously measured at the large scale used in this study
and thus represents new knowledge, nor have implications for aquatic resource
management been considered.
Trace element chemistry, once considered a subdiscipline of geochemistry
(Durum and Haffty 1961), is an important field in its own right. Recent methodological
and analytical advances (Shiller 2003) enable reliable, precise sampling in remote field
locations without the need for clean lab facilities. From a geologic perspective, trace
elements permit examination of the origins (i.e., petrogenesis) of basaltic and igneous
rocks (Hanson 1978). From an applied management perspective, trace element
depositions in calcified structures of aquatic organisms yield critical biological and
ecological information about these species. For example, an emerging research technique
called otolith microchemistry may be used to evaluate natal origins and movement of
fishes by relating trace element concentrations in calcified structures (e.g., otoliths, fin
rays) to water signatures (Zeigler and Whitledge 2010, 2011; Gahagan et al. 2012). If
water chemistry is spatially variable, temporally stable, and proportionally related to
otolith chemistry, trace elements in otoliths function as “natural tags” for assessing fish
environmental history. Thus, documenting spatial and temporal patterns in water
32 chemistry can facilitate discoveries in fisheries research (e.g., spawning locations,
movement rates) that lead to science-based management strategies (e.g., habitat
protection/restoration, harvest regulations) beneficial for biota and humans alike.
Overall, our results enhance understanding of geological influences on water
chemistry in Missouri River reservoirs. In addition, they lay a foundation for research
that necessitates trace element blueprints such as the one established herein. Future
studies examining spatial variability in trace element chemistry on smaller scales (e.g.,
within tributaries and embayments) would permit a higher degree of resolution for water
quality and organismal environmental history research. Moreover, additional research on
stable isotopes (e.g., δ13C, δ18O, 87Sr/86Sr) would expand the scope of geological
knowledge in Missouri River reservoirs and facilitate advancements in aquatic resource
research and management.
Acknowledgements
We thank R. Johnston, C. Hayer, T. Rapp, and J. Mecham of South Dakota State
University for field assistance. We thank B. Hanten and K. Potter of the South Dakota
Department of Game, Fish and Parks and P. Bailey of the North Dakota Game and Fish
Department for technical advice and assistance. We thank A. Shiller and lab members at
the University of Southern Mississippi for providing water sampling kits and measuring
trace element concentrations in water samples. Funding for this project was provided by
the Federal Aid in Sport Fish Restoration program, Project 3M3328, Study 1526,
administered by the South Dakota Department of Game, Fish and Parks, United States
33 Fish & Wildlife Service, South Dakota State University Department of Natural Resource
Management, and South Dakota Agricultural Experiment Station.
34 References
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streams using dialysis in situ – I. Principle and conditions. Water Res. 14:511–
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Winston MR. 2005. Spatiotemporal patterns and changes in Missouri River
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DJ, editors. Biogeochemical cycling of mineral-forming elements. The
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Ridings JW, Crites JW, Hrabik RA, Doyle WJ, Hill TD. 2012. Identifying river of
origin for age-0 Scaphirhynchus sturgeons in the Missouri and Mississippi rivers
using fin ray microchemistry. Can J Fish Aquat Sci. 69:930–941.
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ecological value and vulnerability. Washington (DC): World Resources Institute.
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119.
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differentiate fishes from the Middle Mississippi River, its tributaries, and
floodplain lakes. Hydrobiologia 661:289–302.
36 Tables
Table 2-1. Morphometric and hydrological features of North Dakota and South Dakota
Missouri River reservoirs. Modified from Erickson et al. (2008). Reservoir
Oahe Sharpe Francis Case Lewis and Clark
Oahe Big Bend Fort Randall Gavins Point
1958 1963
1952
1955
28.5
2.2
6.7
0.6
3,621 322
869
145
372
129
172
40
1,263 231
312
97
Feature
Dam Name
Year Closed
3
Gross Volume (km )
Shoreline Length (km)
Reservoir Length (km)
2
Surface Area (km )
6
3
Estimated annual sediment inflow (10 m )
3
Average inflow (m /s)
Exchange rate (years)
24.4
5.3
22.6
3.2
818
1.11
818
0.09
850
0.25
906
0.02
37 Table 2-2. Results of one-way ANOVA testing for spatial variability in water element:Ca
ratios among Missouri River tributaries and sites within reservoirs. A significant P value
(≤ 0.05) indicates water chemistry is spatially variable in the associated water body.
Element Water body
Sr:Ca
Tributaries
Ba:Ca
Mg:Ca
Mn:Ca
Na:Ca
F
F8,6 = 2155.50
P
< 0.01
Oahe tributaries
F5,3 = 73.90
< 0.01
Oahe
F15,13 = 1914.70 < 0.01
Sharpe
F7,6 = 169.28
< 0.01
Francis case
F7,8 = 316.02
< 0.01
Lewis & clark
F6,6 = 1528.10
< 0.01
Tributaries
F8,6 = 7468.80
< 0.01
Oahe tributaries
F5,3 = 1750.80
< 0.01
Oahe
F15,13 = 747.79
< 0.01
Sharpe
F7,6 = 3087.40
< 0.01
Francis case
F7,8 = 158.64
< 0.01
Lewis & clark
F6,6 = 817.86
< 0.01
Tributaries
F8,6 = 5093.40
< 0.01
Oahe tributaries
F5,3 = 1017.20
< 0.01
Oahe
F15,13 = 792.22
< 0.01
Sharpe
F7,6 = 16.68
< 0.01
Francis case
F7,8 = 4529.30
< 0.01
Lewis & clark
F6,6 = 459.41
< 0.01
Tributaries
F8,6 = 32.95
< 0.01
Oahe tributaries
F5,3 = 4723.10
< 0.01
Oahe
F15,13 = 349.26
< 0.01
Sharpe
F7,6 = 703.18
< 0.01
Francis case
F7,8 = 313.41
< 0.01
Lewis & clark
F6,6 = 493.06
< 0.01
Tributaries
F8,6 = 6803.30
< 0.01
Oahe tributaries
F5,3 = 2576.30
< 0.01
Oahe
F15,13 = 2132.50 < 0.01
Sharpe
F7,6 = 142.93
< 0.01
Francis case
F7,8 = 1080.90
< 0.01
Lewis & clark
F6,6 = 1086.77
< 0.01
38 Figures
Figure 2-1. Map of South Dakota surficial geology. Geologically young glacial sediments
predominate east of the Missouri River, whereas older Triassic (Tr), Jurassic (J),
Cretaceous (C), Tertiary (T) sediments exist west of the river. M denotes metamorphic
rocks. Image obtained from South Dakota Department of Environment & Natural
Resources (http://www.sdgs.usd.edu/geologyofsd/geosd.html).
39 Figure 2-2. Sr:Ca and Ba:Ca ratios for tributaries (black boxes) and select
emainstem/embayment sites (gray boxes) in Missouri River reservoirs (Oahe, Sharpe,
Francis Case [FC], and Lewis & Clark [LC]) in North Dakota and South Dakota, USA.
All tributaries and a representative set of mainstem/embayment sites sampled in July
2012 and February 2014 are depicted. Tributaries exhibit spatial variability (i.e., north–
south gradient) in water Sr:Ca and Ba:Ca ratios, whereas mainstem/embayment sites
generally have similar water signatures. Means with the same letter are not significantly
different (ANOVA with Tukey’s HSD test; P < 0.05).
40 Figure 2-3. Mean water Sr:Ca values at 20 collection sites in lakes (a) Oahe, (b) Sharpe,
(c) Francis Case, and (d) Lewis & Clark. Means within reservoirs with the same letter are
not significantly different (ANOVA with Tukey’s HSD test on log10 transformed values;
P < 0.05). Error bars represent SEs. Site codes for Lake Oahe: BVB = Beaver Bay, BVC
= Beaver Creek, CAR = Cannonball River, CRC = Cheyenne River Confluence, CRU =
Cheyenne River Upstream, GRC = Grand River Confluence, GRU = Grand River
Upstream, HAZ = Hazelton, HTR = Heart River, KNR = Knife River, MIN =
Minneconjou Embayment, MOB = Mobridge Main Channel, MRC = Moreau River
Confluence, MRU = Moreau River Upstream, ODF = Oahe Dam Face, OKO = Okobojo
Embayment, SWC = Swan Creek Embayment, WPL = West Pollock Embayment, WWE
= West Whitlocks Embayment, and WWM = West Whitlocks Main Channel. Site codes
for Lake Sharpe: ANT = Antelope Creek, BRC = Bad River Confluence, BRU = Bad
River Upstream, FTG = Fort George, HIP = Hipple Lake, JOE = Joe Creek, NSH = North
Shore, SSB = Sharpe Stilling Basin, and WEB = West Bend. Site codes for Lake Francis
Case: AMC = American Creek, CED = Cedar Shore Main Channel, NBE = North Bay
Embayment, NBM = North Bay Main Channel, PCE = Platte Creek Embayment, PCM =
Platte Creek Main Channel, WRC = White River Confluence, and WRU = White River
Upstream. Site codes for Lewis & Clark Lake: GPD = Gavin’s Point Dam, LCE = Lewis
& Clark Embayment, LCM = Lewis & Clark Main Channel, NRC = Niobrara River
Confluence, NRU = Niobrara River Upstream, SPE = Springfield Embayment, and SPM
= Springfield Main Channel.
41 Figure 2-4. Mean water Sr:Ca values at nine tributaries in Missouri River reservoirs
(Oahe, Sharpe, Francis Case, Lewis & Clark). Means with the same letter are not
significantly different (ANOVA with Tukey’s HSD test on log10 transformed values; P <
0.05). Error bars represent SEs. Site codes are as follows: CAR = Cannonball River,
CHR = Cheyenne River, GRR = Grand River, HTR = Heart River, KNR = Knife R,
MOR = Moreau River, BAR = Bad River, WHR = White River, and NBR = Niobrara R.
42 Figure 2-5. Mean water Ba:Ca values at 20 collection sites in lakes (a) Oahe, (b) Sharpe,
(c) Francis Case, and (d) Lewis & Clark. Means within reservoirs with the same letter are
not significantly different (ANOVA with Tukey’s HSD test on log10 transformed values;
P < 0.05). Error bars represent SEs. Site codes for Lake Oahe: BVB = Beaver Bay, BVC
= Beaver Creek, CAR = Cannonball River, CRC = Cheyenne River Confluence, CRU =
Cheyenne River Upstream, GRC = Grand River Confluence, GRU = Grand River
Upstream, HAZ = Hazelton, HTR = Heart River, KNR = Knife River, MIN =
Minneconjou Embayment, MOB = Mobridge Main Channel, MRC = Moreau River
Confluence, MRU = Moreau River Upstream, ODF = Oahe Dam Face, OKO = Okobojo
Embayment, SWC = Swan Creek Embayment, WPL = West Pollock Embayment, WWE
= West Whitlocks Embayment, and WWM = West Whitlocks Main Channel. Site codes
for Lake Sharpe: ANT = Antelope Creek, BRC = Bad River Confluence, BRU = Bad
River Upstream, FTG = Fort George, HIP = Hipple Lake, JOE = Joe Creek, NSH = North
Shore, SSB = Sharpe Stilling Basin, and WEB = West Bend. Site codes for Lake Francis
Case: AMC = American Creek, CED = Cedar Shore Main Channel, NBE = North Bay
Embayment, NBM = North Bay Main Channel, PCE = Platte Creek Embayment, PCM =
Platte Creek Main Channel, WRC = White River Confluence, and WRU = White River
Upstream. Site codes for Lewis & Clark Lake: GPD = Gavin’s Point Dam, LCE = Lewis
& Clark Embayment, LCM = Lewis & Clark Main Channel, NRC = Niobrara River
Confluence, NRU = Niobrara River Upstream, SPE = Springfield Embayment, and SPM
= Springfield Main Channel.
43 Figure 2-6. Mean water Ba:Ca values at nine tributaries in Missouri River reservoirs
(Oahe, Sharpe, Francis Case, Lewis & Clark). Means with the same letter are not
significantly different (ANOVA with Tukey’s HSD test on log10 transformed values; P <
0.05). Error bars represent SEs. Site codes are as follows: CAR = Cannonball River,
CHR = Cheyenne River, GRR = Grand River, HTR = Heart River, KNR = Knife R,
MOR = Moreau River, BAR = Bard River, WHR = White River, and NBR = Niobrara
River.
44 CHAPTER 3: OTOLITH MICROCHEMISTRY REVEALS NATAL ORIGINS OF
WALLEYES IN MISSOURI RIVER RESERVOIRS
This chapter is being prepared for submission to North American Journal of Fisheries
Management and was co-authored by Mark J. Fincel, Christopher M. Longhenry, and
Brian D. S. Graeb.
45 Abstract
The study of natal origins is important for fisheries management and
conservation. Spawning, rearing, and nursery habitats are vital to sustain populations, but
their identity and location are often unknown or poorly understood. We examined natal
origins of Walleyes Sander vitreus in Missouri River reservoirs, South Dakota, USA,
using otolith microchemistry. Sr:Ca and Ba:Ca ratios at 14 locations in Lakes Oahe,
Sharpe, Francis Case, and Lewis & Clark were spatially heterogeneous and temporally
consistent. Terminal otolith signatures of age-0 Walleyes were temporally matched with
water chemistry and varied in accordance with spatial variability in water chemistry,
representing a blueprint for evaluation of adult natal origins. Seventy-four Walleyes
hatched in Lake Oahe tributaries, representing a relatively large percentage of all adults
(32.31%) and the majority of Lake Oahe adults (77.08%) collected in this study. The
Moreau River and Cannonball River were particularly important natal sites in Lake Oahe,
as were North Shore, West Bend, Platte Creek, and the Lewis & Clark Delta in southern
reservoirs. Overall, otolith microchemistry is an effective, reliable tool for studying natal
origins of Walleyes in Missouri River reservoirs, permitting estimation of relative natal
contribution from tributaries and mainstem/embayment locations. Results from this study
will help inform egg take, habitat conservation, harvest regulations, and other walleye
management activities in Missouri River reservoirs.
46 Introduction
Understanding natal origins is important for management of fisheries throughout
the world. However, knowledge of natal origins is often limited to anecdotal patterns of
site identity and importance. When spatial patterns in natal contribution are unknown,
managers may be unable to accurately prioritize sites for protection and/or restoration.
Natal origins are also pertinent to invasive species management and control (Howe et al.
2013). Reliable information on provenance of invasive fishes is required for targeted
harvest. Despite its importance in fisheries management, knowledge of natal origins is
often fragmentary and would benefit from methods that enable precise measurement of
relative natal contribution.
Walleyes Sander vitreus employ three general life history typologies (Bozek et al.
2011): (1) river resident-river spawning; (2) lake resident-lake spawning; and (3) lake
resident-river spawning. Walleyes in Missouri River reservoirs, especially Lake Oahe,
use the lake resident-river spawning strategy, as do populations in Lake Nipigon
(Dymond 1926), Oneida Lake (Adams and Hankinson 1928), Lake Erie (Wolfert 1963;
Schneider and Leach 1977; Gatt et al. 2003), Lake Ontario (Smith 1892; Bensley 1915),
and Lake Superior (Geiling et al. 1996). Walleyes select spawning sites with coarse
substrate (Scott 1967; Grinstead 1971) to minimize egg siltation during the hatching
period. They also prefer spawning sites with high fetch to increase water and oxygen
flow over eggs (Becker 1983; Martin et al. 2013). A number of diverse habitats may offer
these conditions, including lake tributaries (Pflieger 1997; Kelder and Farrell 2009;
Chalupnicki et al. 2010), flooded marshes (Priegel 1970), riverine portions of reservoirs
(Quist et al. 2004), and riprap on the face of reservoir dams (Grinstead 1971).
47 Understanding Walleye natal origins is important for spawning site protection,
habitat restoration, and conservation planning in Missouri River reservoirs. However,
current knowledge of natal origins in is confined to anecdotal evidence of tributary and
embayment spawning (Mark J. Fincel; South Dakota Department of Game, Fish & Parks
[SDGFP; personal communication). Managers and biologists have ample observational
evidence of natal site identity accumulated through decades of fisheries management
activities, but relative natal contribution is unknown. Quantitative information regarding
the frequency and magnitude of spawning will permit managers to compare the
importance of natal sites and ultimately design science-based management strategies.
Otolith microchemistry is an innovative technique for investigating fish
environmental history. The technique has been used throughout the world to investigate
natal origins (Ruttenberg et al. 2005; Wolff et al. 2012; Rohtla et al. 2014), movement
(Brenkman et al. 2007; Allen et al. 2009), and stock composition (Bickford and Hannigan
2005; Coghlan et al. 2007). Using the proportional relationship between water and otolith
trace element concentrations, otolith microchemistry enables retrospective site
assignment provided water and otolith chemistry are spatially variable and temporally
stable. As such, otolith microchemistry permits confirmation of anecdotal patterns in
natal site identity and importance and estimation of percent natal contribution from
habitats. With this information, managers may be able to prioritize sites for egg take,
habitat protection and improvement, harvest regulations, and other management activities
and thereby advance Walleye management in Missouri River reservoirs.
Our objectives were to assess spatial and temporal patterns in water trace element
chemistry of Missouri River reservoirs, examine relationships between water and otolith
48 chemistry, and evaluate the utility of otolith microchemistry for assessing relative and
site-specific natal contributions. Given geologic heterogeneity in the study area, we
hypothesized that water trace element signatures would be spatially variable, temporally
stable, and positively related to otolith concentrations, rendering otolith microchemistry a
feasible research technique. We predicted that tributaries, particularly those in Lake
Oahe, would be important natal sites and recruitment sources for adult Walleyes.
Methods
Study Area
The Missouri River is the longest river in North America, flowing 3,768 km from
Brower’s Spring, Montana to its confluence with the Mississippi River at St. Louis,
Missouri. It spans 1,371,017 km2, the second-largest riverine drainage area in the United
States, and encompasses 47 tributaries with drainage basins greater than 1000 km2 (Galat
et al. 2005). Much of the Missouri River lies within the semi-arid Great Plains, with half
of the basin receiving less than 41 cm y-1 of precipitation (Galat et al. 2005). Average
July-October water temperatures range from 21.5oC to 27oC in most of the river, but
temperatures below reservoirs are depressed due to hypolimnetic water releases (Galat et
al. 2001). Cropland (37%) and grassland (30%) dominate land use within the basin,
which is minimally developed by humans (9%; Revenga et al. 1998). The South Dakota
portion of the Missouri River is impounded into lakes Oahe, Sharpe, Francis Case, and
Lewis & Clark and is characterized by spatial heterogeneity in surficial geology.
Cenozoic glacial sediments (e.g., Illinois and Wisconsin glacial sediments) predominate
east of the Missouri River, whereas older Mesozoic sediments (e.g., sandstonces, shales,
and clays) exhibit a north-south gradient west of the river.
49 Trace Element Sampling
Water samples (n = 2/site) were collected from tributaries (n = 6), embayments (n
= 7), and a mainstem location (n = 1) in South Dakota Missouri River reservoirs in late
summer 2012 and late winter 2014 to assess spatial and temporal patterns in trace
element concentrations (Figure 3-1). Sampling sites represented known Walleye natal
areas based on recent and historical SDGFP sampling. A syringe filtration method for
sampling in remote field locations (Shiller 2003) and used in previous otolith
microchemistry studies (Zeigler and Whitledge 2010, 2011; Phelps et al. 2012) was
employed. All water samples were collected with nitrile gloves and stored in sealed
coolers (i.e., no light penetration) before filtration. Concentrations of trace elements (i.e..,
strontium [Sr], barium [Ba], magnesium [Mg], manganese [Mn], and sodium [Na]) and
calcium were quantified using high-resolution inductively coupled plasma mass
spectrometry (HR-ICPMS) at the University of Southern Mississippi. These elements are
deposited in otoliths (Campana 1999) and have been used to identify natal habitats in
previous studies (Campana et al. 2000, Brazner et al. 2004). All water elemental
concentrations were converted to molar ratios against calcium (mmol·mol-1) as this
element is a pseudointernal standard (Bickford and Hannigan 2005; Ludsin et al. 2006;
Whitledge et al. 2007).
Fish Sampling
In partnership with SDGFP, age-0 Walleye were collected in late summer 2012
concurrently with water sampling at tributaries (n = 16 individuals), embayments (n = 28
50 individuals), and a mainstem location (n = 2 individuals) using nearshore electrofishing
(Figure 3-1). Total length of the 46 individuals ranged from 76 to 114 mm. If age-0
Walleye and water samples were not collected on the same day due to logistical sampling
constraints, otoliths were ablated in non-terminal locations to ensure temporal matching
with water chemistry (see description below). As with water sampling, juvenile collection
sites represented known Walleye natal areas based on recent and historical SDGFP
sampling.
A total of 243 adult Walleye were collected in 19 embayment and mainstem
locations (n = 10–18/site) in summer 2013 using a variety of methods, including shortterm gill nets, deepwater gillnets, and angler donations. The total length of each
individual was recorded and ranged from 190 to 756 mm. Adults ranged from age-2 to
age-11 with the majority (68.31%) age-2 or age-3, nearly one-quarter (23.87%) age-4 or
age-5, and older age classes each represented by 1–8 individuals.
Otolith Microchemistry
Otolith microchemistry is a sensitive technique that can be used to evaluate fish
natal origins. Otoliths are paired calcified structures that fishes use for hearing and
balance. They permanently deposit trace elements in proportion to water column
concentrations. Combined with otolith biochronological properties, elemental
accumulation permits retrospective assessment of environmental history provided water
signatures are spatially heterogeneous and temporally constant.
Otolith microchemistry requires an extensive protocol to prevent contamination
and erroneous evaluation of natality, movement, and entrainment. In this study, Walleyes
51 were sacrificed immediately after collection, placed in individual labeled bags, and stored
on ice in sealed coolers until same-day otolith extraction in a laboratory. Left and right
sagittal otoliths from each individual were removed using plastic forceps triple-washed in
nitric acid (Campana et al. 2000; Brazner et al. 2004). The otolith with the most welldefined annuli was used for ageing by three independent readers (correspondence >
90%). In preparation for trace element analysis, otoliths were triple-rinsed in distilled,
ultrapure water; air-dried for a minimum of 24 hrs; and stored in acid-washed 2-mL
polypropylene microcentrifuge tubes (Zeigler and Whitledge 2010). After initial
cleaning, adult otoliths were embedded in Epo-Fix epoxy and sectioned in the transverse
plane using a low-speed Isomet diamond saw (Buehler, Lake Bluff, Illinois). Each
section included the otolith core, and contamination was prevented by cleaning the saw
blade with aluminum oxide lapping film after each section. Age-0 otoliths were not
sectioned due to small size and fragility but rather placed in thermoplastic glue and
ground in the sagittal plane. All otoliths were sanded evenly and polished with 3M Wet
or Dry sandpaper (400 grit) and aluminum oxide lapping film, mounted on acid-washed
petrographic slides (Donohoe and Zimmerman 2010), triple-sonicated in ultrapure water
before trace element analysis, and dried in a Class 100 laminar flow hood for 24 hr.
Trace element (i.e.., 88Sr, 86Sr, 138Ba, 137Ba, 24Mg, 55Mn, and 23Na) concentrations
were quantified with laser ablation inductively coupled plasma mass spectrometry (LAICPMS) at the nterdisciplinary Center for Inductively Coupled Plasma Mass
Spectrometry at the University of California – Davis. An Agilent Technologies 7500a
quadrupole ICP-MS coupled to a New Wave Research UP-213-nm laser with He as the
carrier gas was used for laser ablation analysis. Only 88Sr and 137Ba were used to assess
52 Walleye natal origins as they were the only signatures that were spatially variable and
temporally stable (Pracheil et al. 2014). Laser parameters were 70 % energy, 10 Hz, 40
um spot size, 25 sec dwell time, 50 sec acquisition and 25 sec background. USGS
synthetic glass standard GSE-1G was used as the calibration standard, and two additional
reference standards (GSD-G1 and MACS-3) were used as quality controls for verification
of instrument accuracy and precision. Each standard was ablated in three to five locations
after every four samples to adjust for possible instrument drift. Mean limits of detection,
calculated as mean blank values plus three standard deviations (Wells et al. 2003), were
0.01 and 0.07 for 88Sr and 137Ba, respectively; concentrations of these elements in all
otoliths were well above detection limits. Specialized computer software (Glitter 4.4;
GEMOC CSIRO, Macquarie Research Ltd., Macquarie University, Sydney, Australia)
was used for data reduction. Otolith signatures were background subtracted, 43Ca was
used as an internal standard (Bickford and Hannigan 2005; Ludsin et al. 2006; Whitledge
et al. 2007), and final trace element concentrations were obtained by matching 43Ca
counts to the sample CaO concentrations obtained independently by electron microprobe.
All otolith microchemistry data are reported as element:Ca ratios (μmol·mol-1). Only
signatures that were both spatially variable and temporally consistent were used to assess
Walleye natal origins.
All otoliths were ablated using spot analyses. For each spot, a 15-s laser warm-up
time was followed by a 20-s dwell time during which the sample was ablated. The
integration time for all elements (0.01 s for 43Ca, 0.05 s for 88Sr and 137Ba) was repeated
throughout the 20-s dwell time. Following each ablation, there was a 95-sec washout
time. Natal signatures of age-0 and adult Walleyes were quantified by ablating otoliths in
53 the core (Ruttenberg et al. 2005). Adult Walleye otoliths were ablated at each annuli and
inter-annual areas proceeding outward from cores to evaluate site occupancy from
hatching to collection.
Age-0 otoliths were also ablated to characterize recent environmental history in a
manner that ensured temporal matching with water chemistry. When otolith and water
samples were collected at the same time, signatures were matched by ablating otoliths at
terminal edges, which reflect recently occupied environments (Zeigler and Whitledge
2010, 2011). Hereafter these signatures are referred to as “mean terminal.” When waterage-0 synchronization was not possible, age-0 individuals were invariably collected after
water samples, which permitted laser ablation at non-terminal otolith locations that
coincided with the time of water sampling. To quantify these “adjusted mean terminal”
signatures, water and otolith chemistry were synchronized by quantifying the number of
days between water and age-0 sampling and ablating otoliths at locations an equivalent
number of daily rings from otolith edges. Temporal matching between otolith and water
chemistry ensured accurate spatial blueprinting (e.g., otolith signatures represented
known capture locations). In addition, age-0 otolith signatures were generally stable from
core to edge, verifying our supposition that juveniles remained in natal environments
until capture. Overall, our efforts to ensure temporal matching between water and age-0
signatures mirror those of recent studies (Zeigler and Whitledge 2010, 2011), in contrast
to previous investigations that were either unclear about temporal matching (Wells et al.
2003, Whitledge et al. 2007) or did not involve water sampling (Brazner et al. 2004,
Cairns et al. 2004, Downs et al. 2006). Although explicit measurement of water
chemistry is not required for reliable otolith microchemistry, particularly in large river
54 systems such as the Missouri River and Eleven Point River (Bickford and Hannigan
2005) in which water signatures are principally influenced by stable geology and
weathering (Gibbs 1970), we quantified water chemistry and temporally matched it with
age-0 otolith chemistry to ensure reliable, accurate results.
Statistical Analysis
Spatial (i.e., site-level) and temporal (i.e., annual) patterns in water chemistry
were assessed using two-way analysis of variance (ANOVA) with Tukey’s Honestly
Significant Difference (HSD) test for multiple comparisons on log10-transformed water
data to ensure normality and equal variances (Gahagan et al. 2012). One-way ANOVA
with Tukey’s HSD test was used to evaluate differences in water chemistry among
reservoirs and variability in age-0 natal signatures among collection sites. Water-otolith
trace element relationships were evaluated with least-squares linear regression (Munro et
al. 2005; Zeigler and Whitledge 2010; Phelps et al. 2012) using water and age-0 terminal
(i.e., mean terminal, adjusted mean terminal) otolith signatures from the 14 sites where
both samples were collected. Statistical significance for all analyses was set at α ≤ 0.05.
The accuracy with which age-0 Walleye could be classified to known collection
site types (i.e., tributary, embayment, mainstem) and individual sites sites based on
bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures was evaluated using K-sample nearest
neighbor discriminant analysis. Used in previous microchemistry research (Bickford and
Hannigan 2005), this nonparametric method allows for reliable classification when
otolith data do not meet parametric assumptions (i.e., multivariate normality, equal
variance-covariance matrices). This procedure assigns age-0 individuals to natal sites to
55 which the majority of their k nearest neighbors belong (Johnson 1998). The accuracy of
different models (i.e., k = 2–8) was evaluated using a leave-one-out jackknife procedure,
and the model with the lowest error rate (k = 2) was used to classify adults to natal sites
using the known-origin data set (Ruttenberg et al. 2005). Juvenile Walleyes were used to
develop the discriminant model under the notion that they would reflect site-specific
signatures of capture locations unlike mobile adults, a supposition that was verified
through comparison of core and edge signatures. Adult natal origins were summarized as
percentage contributions from different tributary, embayment, and mainstem sites.
Results
Water Chemistry
Element:Ca ratios were spatially heterogeneous at two levels (i.e., all sites,
tributaries; Table 3-1) in the same way each year for Sr:Ca (F1,179 = 69.89, P = 0.10) and
Ba:Ca (F1,179 = 102.62, P = 0.49) and different ways annually for Mg:Ca (F1,179 = 20.69,
P < 0.01), Mn:Ca (F1,91 = 32.95, P < 0.01), and Na:Ca (F1,103 = 45.50, P < 0.01).
Concentrations of temporally stable signatures useful for otolith microchemistry (i.e.,
Sr:Ca, Ba:Ca) were also heterogeneous among reservoirs (Sr:Ca: F3,67 = 5.39, P < 0.01;
Ba:Ca: F3,67 = 7.30) and among the specific sites where age-0 Walleye were collected
(Sr:Ca: F13,14 = 919.16, P < 0.01; Ba:Ca: F13,14 = 814.61, P < 0.01; Figure 3-2) in late
summer 2012. Tributaries on the western side of the Missouri River exhibited a
decreasing north-south Sr:Ca gradient consistent with latitudinal variation in surficial
geology. Mainstem sites on the eastern side of the river generally had more consistent
56 Sr:Ca ratios. Ba:Ca ratios of western tributaries followed the opposite north-south
pattern, increasing moving downstream, but eastern mainstem sites were relatively
homogenous.
Otolith Chemistry
Terminal (i.e., mean terminal, adjusted mean terminal) otolith signatures of age-0
Walleyes were heterogeneous for Sr:Ca and Ba:Ca (Figure 3-3) in accordance with
temporal stability (Sr:Ca: F1,57 = 69.89, P = 0.10; Ba:Ca: F1,57 = 102.62, P = 0.49) and
spatial variability (Table 3-1) in water chemistry in Missouri River reservoirs. Waterotolith relationships were positive and proportional for Sr:Ca (R2 = 0.71, P < 0.01; Figure
3-4a) and Ba:Ca (R2 = 0.40, P = 0.02; Figure 3-4b). Age-0 Walleye were classified with
high accuracy to site types (95% mean accuracy) and individual sites (80% mean
accuracy; Table 3-2) using k-sample nearest neighbor discriminant analysis. Accuracy
was higher for tributaries (100%) than embayments (86%) and 100 percent for the single
mainstem site (i.e., Ft. George). Site-specific terminal otolith signatures of age-0
Walleyes constituted a chemical blueprint for examination of adult natal origins.
Natal Contribution
Spatial variability in Sr:Ca and Ba:Ca signatures enabled accurate natal
site identification of age-0 and adult Walleyes. Seventy four Walleyes hatched in Lake
Oahe tributaries from 2002–2012, representing a relatively large percentage of all adults
(32.31%) and the majority of Lake Oahe adults (77.08%) collected in this study. Natal
contributions of tributaries, including the Moreau River (38.54%, n = 37) and Cannonball
57 River (18.75%, n = 18), were large compared to all mainstem/emaybment sites combined
(22.92%, n = 22; Table 3). Natal contribution of the Moreau River was 57.1 percent (n =
16) for Lake Oahe individuals and 39.5 percent (n = 17) overall in 2009 during a prairie
flood, when 85.7 percent (n = 24) of Lake Oahe individuals and 58.1 percent (n = 25) of
all Walleyes hatched in tributaries. Natal contribution of tributaries was also appreciable
in 2010 (47.7%, n = 42) and 2011 (47.7%, n = 31). Across years, it declined
progressively moving downstream from Lake Oahe (48.03%, n = 46) to Lake Sharpe
(32.69%, n = 17), Lake Francis Case (25.00%, n = 10), and Lewis & Clark Lake
(22.50%, n = 9). Beaver Bay was an important natal site for Walleyes that did not hatch
in tributaries.
Approximately one-third (32.31%, n = 17) of adult Walleyes collected in Lake
Sharpe hatched in tributaries from 2002-2012. Of these, only 23.53% (n = 4) originated
in the Bad River (i.e., Lake Sharpe), whereas 76.47% (n = 13) hatched in Lake Oahe
tributaries (Table 3). One-fourth of adult Walleyes collected in Lake Francis Case
hatched in the Bad River or the White River from 2002-2012 (Table 3). Unlike other
reservoirs, no individuals from Lake Francis Case originated in Lake Oahe tributaries.
Nine adult Walleyes collected in Lewis & Clark Lake hatched in Lake Oahe tributaries
from 2002-2012. The Lewis & Clark Delta was an important natal site, contributing
57.50% of adult Walleyes in that reservoir (Table 3).
Discussion
Otolith microchemistry has powerful applications for the study of fish
environmental history. Research throughout the world has spanned an array of topics,
58 including reproductive ecology (Ruttenberg et al. 2005; Wolff et al. 2012; Rohtla et al.
2014), movement (Brenkman et al. 2007; Allen et al. 2009), entrainment (Shima and
Swearer 2009), stock discrimination (Bronte et al. 1996; Bickford and Hannigan 2005),
population marking (Kennedy et al. 2000), habitat fingerprinting (Brazner et al. 2004),
and invasion biology of nonnative species (Munro et al. 2005; Whitledge et al. 2007).
Our results contribute to this growing body of literature as otolith microchemistry was an
effective, reliable tool for studying Walleye natal origins in Missouri River reservoirs and
tributaries. To our knowledge, this research was only the third otolith microchemistry
study to examine Walleye natal origins and the first in the Missouri River. Spatial
variability and temporal stability (Bickford and Hannigan 2005) in water trace element
chemistry and proportional water-otolith relationships laid a foundation for reliable,
informative microchemical analysis. Terminal otolith Sr:Ca and Ba:Ca ratios of age-0
Walleyes were spatially heterogeneous in accordance with water chemistry, particularly
for Missouri River tributaries, and thus represented elemental fingerprints adult site
assignment. However, Mg:Ca, Mn:Ca, and Na:Ca did not reflect water chemistry and
were thus unreliable signatures. Mg and Na are loosely bound in the CaCO3 matrix and
thus more prone to chemical alteration than tightly bound elements such as Sr and Ba
(Milton and Chenery 1998, Proctor and Thresher 1998, Hedges et al. 2004). Our results
confirm previous research that Mn concentrations are generally low (Wells et al. 2003)
and unstable (Wetzel 1983) in freshwater compared to marine environments where this
element is a highly effective chemical tracer (Pracheil et al. 2014). Lower utility in
freshwater may reflect intricate metabolic processes (Gibson-Reinemer et al. 2009) or
growth-related biogeochemical changes (Limburg et al. 2010). Overall, we expect Sr:Ca
59 and Ba:Ca to be most effective for future otolith microchemistry research in Missouri
River reservoirs.
Site-specific trace element signatures of Missouri River impoundments were
distinct and represented a powerful tool for delineating natal origins of Walleyes,
confirming previous studies on this species (Bickford and Hannigan 2005), other percids
(Brazner et al. 2004), salmonids (Martin et al. 2012), centrarchids (Zeigler and Whitledge
2011), temperate basses (Zeigler and Whitledge 2010), and acipenserids (Phelps et al.
2012). Natal site classification using k-sample nearest neighbor discriminant analysis was
highly accurate for tributaries, particularly those in Lake Oahe, reflecting bivariate
distinctness in water chemistry resulting from heterogeneous surficial and bedrock
geology. Classification was also accurate for embayments and mainstem locations,
indicating Sr:Ca and Ba:Ca are reliable signatures for natal site identification. We
recommend future researchers expand our analysis by evaluating additional trace element
and isotopic signatures as univariate and multivariate tracers of natal origins in Missouri
River reservoirs, particularly in mainstem locations.
A large portion of adult Walleyes and the majority of tributary Walleyes collected
in this study hatched in Lake Oahe tributaries. Unique tributary trace element signatures
were detectable in otoliths of adult Walleyes and revealed the importance and relative
contributions of natal rivers. The Moreau River and Cannonball River had the highest
percent natal contributions, suggesting these rivers are particularly important for
spawning and thereby confirming anecdotal evidence. This finding is especially
significant for the Cannonball River, where a proposed damming project may have
negative consequences on spawning Walleyes and downstream populations (Mark Fincel,
60 SDGFP, personal communication). Walleye natal contribution was also high in the Lewis
& Clark Delta, a refuge environment during floods and non-flood periods (Carlson et al.
2015) where walleye spawning has been observed (Graeb et al. 2009) and species-rich
fish communities have been linked to high-quality reproductive and rearing habitats
(Kaemingk et al. 2007). As such, otolith microchemistry is an auspicious tool for
examining the contribution of this region to spawning and recruitment of Walleye and
other species. Moreover, it will empower decision makers to implement science-based
management strategies.
Our results have diverse management applications in Missouri River
impoundments and supplement a burgeoning body of literature that demonstrates the
utility of otolith microchemistry for fisheries management throughout the world. For
example, we advise managers to focus egg take in sites where high numbers of walleye
reproduce to streamline operations, protect walleye in areas with fewer spawners, and
promote spatially extensive reproduction. For habitat conservation, we suggest managers
protect natal sites from sedimentation, shoreline erosion, damming, pollution, and other
stressors to promote riparian functions and services (e.g., sediment control, nutrient
cycling, flood control) at impoundment and riverscape (Fausch et al. 2002) scales. For
habitat restoration, we recommend managers enhance physical and chemical habitats in
degraded natal areas by dredging sediments, stabilizing shorelines, planting riparian
buffers, and/or mitigating point-source pollution. We advise managers to educate and
incentivize landowners throughout the riverscape to minimize nutrient and sediment
delivery into reservoirs. We also suggest they ensure water level manipulations maintain
or improve (rather than degrade) walleye spawning and nursery habitats. Managers can
61 expect tributary natal contribution to be highest during wet years, which informs site
prioritization for management activities (e.g., egg take, harvest regulations) and indicates
tributary habitat conservation is particularly important during these time periods.
However, in all reservoirs but Lake Oahe the majority of adult walleyes spawned in nontributary habitats. It is important for managers to balance the reproductive importance of
these environments against that of tributaries. We recommend managers combine
anecdotal information on spawning site identity with future research (e.g., telemetry,
otolith microchemistry with novel signatures) to quantify the relative significance of nontributary spawning sites. Moreover, our analysis focused on spawning, which does not
necessarily translate into recruitment, perhaps the most meaningful dynamic rate function
for managers. Thus, we recommend managers use predictive modeling based on existing
and future population data to link spawning with recruitment. This will enable managers
to identify natal sites that function as “recruitment hotspots” and design corresponding
management strategies (e.g., minimum size limits) that protect recruits in these areas.
Otolith microchemistry is a precise, high-resolution technique that confirmed
anecdotal patterns in Walleye natal site identity and importance. It also permitted
estimation of relative natal contribution from Missouri River tributaries and embayment
locations. This corroborates a growing literature on the utility of chemical signatures as
biological tracers of fish stocks (Campana 1999; Campana et al. 2000), particularly in the
context of natal origins (Brazner et al. 2004; Zeigler and Whitledge 2011). However,
contemporary Walleye otolith microchemistry research is limited to frequentist (Bickford
and Hannigan 2005) and Bayesian (Pflugeisen and Calder 2012) stock discrimination
studies. Natal origins information revealed in this study represents a new avenue in
62 Missouri River fisheries research, one that is germane for natural resource agencies
responsible for managing Walleye populations. Moreover, our results add to a growing
body of literature validating the efficacy of otolith microchemistry for fisheries
management throughout the world (Campana et al. 2000, Ruttenberg et al. 2005; Rohtla
et al. 2014).
We recognize anecdotal evidence indicates that Walleyes are not exclusively lake
resident-river spawners in Missouri River reservoirs; mainstem spawning has also been
documented. This represents a limitation of the present study as spatial variability in
water and otolith Sr:Ca and Ba:Ca signatures was highest in tributaries and embayments
and considerably lower elsewhere. Reclassification to mainstem sites was largely
inaccurate. However, this creates an opportunity for future research to identify additional
chemical tracers (e.g., δ13C, δ18O, δ2H, 87Sr/86Sr; Ziegler and Whitledge 2010, 2011;
Rohtla et al. 2014) that vary among mainstem sites and permit accurate reclassification.
These high-resolution tracers would enable researchers to quantify chemical variability
within tributaries and embayments, identify site-specific residence (e.g., tributary
headwaters versus confluence) of Walleye and other Missouri River fishes, and evaluate
natal origins and movements at fine scales. Such advancements would improve Missouri
River Walleye management in particular and fisheries conservation in general through
enhanced management applicability and agency relevance of otolith microchemistry.
Acknowledgments
We thank R. Johnston, C. Hayer, T. Rapp, and J. Mecham of South Dakota State
University for field assistance. We thank B. Hanten, H. Meyer, K. Potter, G. Knecht, J.
63 Sorensen, T. Sorensen, R. Trible, B. Larson of the South Dakota Department of Game,
Fish and Parks and P. Bailey of the North Dakota Game and Fish Department for
technical advice and assistance. We thank A. Shiller and lab members at the University of
Southern Mississippi for providing water sampling kits and measuring trace element
concentrations in water samples. We thank G. Barford, J. Commisso, and J. Glessner at
the University of California–Davis for assistance with otolith microchemistry analyses.
Funding for this project was provided by the Federal Aid in Sport Fish Restoration
program, Project 3M3328, Study 1526, administered by the South Dakota Department of
Game, Fish and Parks, United States Fish & Wildlife Service, South Dakota State
University Department of Natural Resource Management, and South Dakota Agricultural
Experiment Station.
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72 Tables
Table 3-1. Results of one-way ANOVA testing for spatial variability in water element:Ca
ratios among Missouri River tributaries and sites within reservoirs. A significant P value
(≤0.05) indicates water chemistry is spatially variable in the associated water body.
Element Water body
Sr:Ca
Tributaries
Ba:Ca
Mg:Ca
Mn:Ca
Na:Ca
F
F8,6 = 2155.50
P
< 0.01
Oahe tributaries
F5,3 = 73.90
< 0.01
Oahe
F15,13 = 1914.70 < 0.01
Sharpe
F7,6 = 169.28
< 0.01
Francis case
F7,8 = 316.02
< 0.01
Lewis & clark
F6,6 = 1528.10
< 0.01
Tributaries
F8,6 = 7468.80
< 0.01
Oahe tributaries
F5,3 = 1750.80
< 0.01
Oahe
F15,13 = 747.79 < 0.01
Sharpe
F7,6 = 3087.40
< 0.01
Francis case
F7,8 = 158.64
< 0.01
Lewis & clark
F6,6 = 817.86
< 0.01
Tributaries
F8,6 = 5093.40
< 0.01
Oahe tributaries
F5,3 = 1017.20
< 0.01
Oahe
F15,13 = 792.22 < 0.01
Sharpe
F7,6 = 16.68
< 0.01
Francis case
F7,8 = 4529.30
< 0.01
Lewis & clark
F6,6 = 459.41
< 0.01
Tributaries
F8,6 = 32.95
< 0.01
Oahe tributaries
F5,3 = 4723.10
< 0.01
Oahe
F15,13 = 349.26 < 0.01
Sharpe
F7,6 = 703.18
< 0.01
Francis case
F7,8 = 313.41
< 0.01
Lewis & clark
F6,6 = 493.06
< 0.01
Tributaries
F8,6 = 6803.30
< 0.01
Oahe tributaries
F5,3 = 2576.30
< 0.01
Oahe
F15,13 = 2132.50 < 0.01
Sharpe
F7,6 = 142.93
< 0.01
Francis case
F7,8 = 1080.90
< 0.01
Lewis & clark
F6,6 = 1086.77
< 0.01
73 Table 3-2. Results of k-sample nearest neighbor discriminant analysis with leave-one-out
jackknife cross-validation using Sr:Ca and Ba:Ca ratios of age-0 Walleye. Data include
the number known to have come from site types and individual sites (Known), the
number assigned to those locations (Assigned), the percent deviation of Assigned from
Known (Deviation), and the percentage of individuals classified to known locations
(Accuracy). Signatures were measured at otolith locations synchronized with water
sample collection to ensure reliable spatial blueprinting.
Natal origin
Site type
Tributary
Embayment
Mainstem
Site
Bad R.
Beaver Bay
Cannonball R.
Cheyenne R.
Ft. George
Heart R.
Knife R.
Minneconjou
Moreau R.
N. Bay
Platte Creek
Springfield
W. Pollock
W. Whitlocks
Known (number)
Assigned (number)
Deviation (%)
Accuracy (% of known)
16
28
2
19
26
3
19
-7
50
100
86
100
Mean: 95
4
2
2
4
2
2
2
4
2
8
4
2
3
5
7
2
2
3
3
3
2
4
2
8
3
3
1
3
75
0
0
-25
50
50
0
0
0
0
-25
50
-67
-40
100
100
100
75
100
100
100
75
100
63
50
100
33
20
Mean: 80
74 Table 3-3. Percent (number) natal contribution of tributaries and mainstem/embayment
locations to adult Walleye populations in Missouri River reservoirs, North Dakota and
South Dakota, USA, grouped by collection reservoir and site. FC and LC denote Lake
Francis Case and Lewis & Clark Lake, respectively.
Tributaries
Oahe – Cannonball R.
Oahe – Grand R.
Oahe – Heart R.
Oahe – Knife R.
Oahe – Moreau R.
Sharpe – Bad R.
FC – White R.
Mainstem/Embayment Sites
Oahe – Beaver
Oahe – Minneconjou
Oahe – W. Pollock
Oahe – W. Whitlocks
Sharpe – Ft. George
Sharpe – N. Shore
Sharpe – Stilling Basin
Sharpe – W. Bend
FC – American Creek Embayment
FC – Chamberlain Area
FC – Stilling Basin
FC – Platte Creek
LC – GPD
LC – Tailrace
LC – Delta
Oahe
Sharpe
FC
LC
18.75 (18)
5.21 (5)
6.25 (6)
8.33 (8)
38.54 (37)
---
7.69 (4)
1.92 (1)
1.92 (1)
5.77 (3)
7.69 (4)
7.69 (4)
--
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
7.50 (3)
17.50 (7)
12.50 (5)
2.50 (1)
2.50 (1)
0.00 (0)
5.00 (2)
0.00 (0)
0.00 (0)
8.33 (8)
1.04 (1)
6.25 (6)
7.29 (7)
------------
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
19.23 (10)
28.85 (15)
3.85 (2)
15.38 (8)
--------
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
5.00 (2)
7.50 (3)
10.00 (4)
2.50 (1)
15.00 (6)
12.50 (5)
22.50 (9)
----
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
10.00 (4)
0.00 (0)
2.50 (1)
0.00 (0)
0.00 (0)
0.00 (0)
0.00 (0)
5.00 (2)
2.50 (1)
57.50 (23)
75 Figures
Figure 3-1. Water and walleye sampling locations in Missouri River reservoirs. All sites
were sampled for water trace element concentrations using a syringe-filtration protocol
described in Shiller (2003). Shapes denote locations where fish of different ages were
sampled: right-angle triangles (age-0), upright triangles (adult), diamonds (both).
76 Figure 3-2. Mean water (a) Sr:Ca and (b) Ba:Ca value at the 14 sites where both water
and age-0 Walleyes were collected in late summer 2012. Means with the same letter are
not significantly different (ANOVA with Tukey’s HSD test on log10 transformed values;
P ≤ 0.05). Error bars represent 1 SEM.
77 Figure 3-3. Terminal otolith (a) Sr:Ca and (b) Ba:Ca ratios of age-0 Walleyes at the 14
sites where both water and age-0 Walleyes were collected in late summer 2012.
Signatures represent mean terminal or adjusted mean terminal concentrations that are
temporally matched with water sampling. Means with the same letter are not significantly
different (ANOVA with Tukey’s HSD test on log10 transformed values; P ≤ 0.05). Error
bars represent 1 SEM.
78 Figure 3-4. Linear regression of age-0 Walleye terminal otolith (a) Sr:Ca and (b) Ba:Ca
signatures on water ratios at collection sites lakes Oahe, Sharpe, Francis Case, and Lewis
& Clark. Fish and water sampling occurred in late summer 2012. Error bars represent 1
SEM.
79 CHAPTER 4: THE MISSOURI RIVERSCAPE: OTOLITH MICROCHEMISTRY
FACILITATES ECOSYSTEM-BASED FISHERIES MANAGEMENT IN NORTH
AMERICA’S LONGEST RIVER
This chapter is being prepared for submission to Ecosphere and was co-authored by
Mark J. Fincel and Brian D. S. Graeb.
80 Abstract
Ecosystem-scale processes shape the structure and function of riverscapes. Fish
environmental history is a fundamental yet often poorly understood topic that highlights
the importance of provenance, population connectivity, and other ecosystem-level
processes for fisheries research and management throughout the world. In response to a
catastrophic flood in 2011, we examined movement and entrainment of walleye (Sander
vitreus) in Missouri River reservoirs, North Dakota and South Dakota, USA, using otolith
microchemistry. Age-0 walleye were classified to known natal locations with high
accuracy (75–87%) based on natal Sr:Ca and Ba:Ca signatures that were effective
elemental fingerprints for examining adult movement and entrainment. Walleye
movement within and among Missouri River reservoirs was dominated by downstream
passage and site residency before, during, and after the flood. Upstream movement was
comparatively uncommon, especially during the disturbance. The percentage of entrained
walleye progressively increased moving downstream from Lake Sharpe (18.87%) to Lake
Francis Case (24.39%) to Lewis & Clark Lake (34.15%). In addition, entrainment was
magnified during the flood as nearly two-thirds (64.71%) of entrained walleye moved
through dams during the disturbance. Intra-reservoir movement and entrainment play a
vital role in structuring walleye populations in Missouri River reservoirs. Otolith
microchemistry represents a predictive tool to forecast these aspects of environmental
history in diverse hydrological regimes. It provides insight on population connectivity
and illustrates the significance of large-scale, multi-state walleye management. This
riverscape paradigm will be important for fisheries conservation in the Missouri River in
the present and future.
81 Introduction
Research investigating the spatial and temporal distribution of fishes is critical for
management of fisheries throughout the world (Lucas and Baras 2000, Pelicice and
Agostinho 2008, Baumgartner et al. 2014). For example, knowledge of spawning location
and timing permits prioritization of nursery areas for monitoring and protection
(Landsman et al. 2011). Understanding movements to and from stocking sites enables
evaluation and enhancement of rehabilitation programs (Fielder 2002). Moreover,
invasive species management depends on reliable knowledge of spatial ecology (Christie
and Goddard 2003).
Knowledge of movement is critical for understanding spatial and temporal
distributions of fishes and ultimately managing populations (Lucas and Baras 2000).
Movement research to date has relied heavily on mark-recapture and radiotelemetry
(Landsman et al. 2011). Walleye (Sander vitreus) is ecologically and economically
important species in North America. The species makes long-range movements for
spawning and foraging in rivers (Eschmeyer 1950, Rawson 1957), inland lakes (Diana et
al. 1990), and the Great Lakes (Ferguson & Derksen 1971). However, the majority of
walleye movement studies have explored select topics (e.g., reproductive biology,
stocking, and habitat use; Crowe 1962) in lacustrine systems (e.g., Great Lakes; Wang et
al. 2007) using mark-recapture and radiotelemetry. Knowledge of fish movement would
be enhanced through implementation of new and more powerful technologies, including
biologgers, acoustic telemetry, hydroacoustics/sonar, and otolith microchemistry/isotope
analysis (Landsman et al. 2011).
Fish entrainment occurs when individuals are displaced from reservoirs to
82 downstream waters through turbines or other water release structures (Martins et al.
2013), potentially altering population dynamics, connectivity, and resilience (via
mortality; Barnthouse et al. 1983; Stevens et al. 1985). Moreover, entrainment caused by
stochastic disturbances (e.g., floods) is unpredictable in the absence of well-defined
relationships between water discharge and fish passage. Understanding how entrainment
relates to water discharge is imperative for predicting effects of disturbances on source
and recipient fish populations and developing corresponding management strategies
(Grimaldo et al. 2009).
Catastrophic flooding occurred in the Missouri River in 2011, damaging
residential, corporate, industrial, agricultural, and transportation infrastructure (Grigg et
al. 2011) and causing billions of dollars in expenses. Ecological effects of the flood are
largely unknown and warrant further research. Entrainment of walleye through mainstem
Missouri River dams is known to occur under elevated and normal discharge conditions
(Smith and Brown 2002), but the magnitude and extent of fish passage have not been
evaluated. Quantifying these relationships would facilitate development of predictive
management strategies under diverse hydrological regimes.
We examined entrainment of walleye in Missouri River reservoirs in response to
catastrophic flooding in 2011. Our objectives were to assess spatial and temporal patterns
in trace element chemistry of Missouri River reservoirs and evaluate the utility of otolith
microchemistry as a technique to study walleye entrainment. We hypothesized that water
trace element signatures would be spatially variable, temporally stable, and positively
related to otolith concentrations, rendering otolith microchemistry technique reliable
environmental history tool. We predicted that adult walleye survived entrainment and
83 inter-reservoir movement occurred under normal hydrological conditions but was
magnified during the flood.
Methods
Study Area
The Missouri River is the longest river in North America, flowing 3,768 km and
draining 1,371,017 km2 through a network of 47 large tributaries (i.e.., drainage basin >
1000 km2; Galat et al. 2005). Land cover within the basin is primarily cropland (37%)
and grassland (30%) with minimal human development (9%; Revenga et al. 1998). The
South Dakota portion of the Missouri River is impounded into lakes Oahe, Sharpe,
Francis Case, and Lewis & Clark and is characterized by spatial heterogeneity in surficial
geology. Cenozoic sediments (e.g., Illinois and Wisconsin glacial sediments)
predominate east of the Missouri River, whereas sediments are largely Mesozoic (e.g.,
sandstonces, shales, and clays) west of the river.
Trace Element Sampling
We used a syringe filtration method (Shiller 2003) employed in previous otolith
microchemistry studies (Zeigler and Whitledge 2010, 2011; Phelps et al. 2012) to sample
trace element concentrations from 57 locations in Missouri River reservoirs in late
summer 2012 and late winter 2014 (Figure 1). Surficial and bedrock geology are
temporally stable throughout the four impoundments and likely cause consistent water
element:Ca ratios (Bickford and Hannigan 2005), but sampled water across years to
verify that water chemistry was temporally stable. Sampling sites included main
84 channels, embayments, stilling basins, tailraces, and free-flowing tributaries. Trace
element (i.e.., strontium [88Sr, 86Sr], barium [138Ba, 137Ba], magnesium [24Mg], manganese
[55Mn], and sodium [23Na]) and calcium (43Ca) concentrations were quantified using
high-resolution inductively coupled plasma mass spectrometry (HR-ICPMS) at the
University of Southern Mississippi. These elements accumulate in otoliths and have been
used in previous otolith microchemistry studies (Campana et al. 2000, Brazner et al.
2004).
Fish Sampling
We collected 46 age-0 walleye during their first growing season in late summer
2012 concurrently with water sampling in eight tributaries (n = 16 individuals), seven
embayments (n = 28 individuals), and two mainstem location (n = 2 individuals) using
nearshore electrofishing (Figure 1). We selected juvenile collection sites because they
were known natal areas and widely distributed throughout the four impoundments as
needed for large-scale analysis of movement and entrainment. Juvenile total length
ranged from 76–114 mm. If age-0 walleye and water samples were not collected on the
same day due to logistical sampling constraints, otoliths were ablated in non-terminal
locations to ensure temporal matching with water chemistry (see description below).
We collected 228 adult walleye in 19 embayment and mainstem locations (n =
10–18/site) in summer 2013 using electrofishing and short- and long-term shallow and
deep water gill nets. Adults ranged from 190–756 mm and age-2 to age-11 with the
majority (68.31%) age-2 or age-3, nearly one-quarter (23.87%) age-4 or age-5, and older
age classes each represented by 1–8 individuals.
85 Otolith Microchemistry
We used an extensive protocol to prevent contamination and erroneous evaluation
of movement and entrainment. We sacrificed walleye immediately after collection,
placed them in individual labeled bags, and stored them on ice in sealed coolers before
same-day otolith extraction in a laboratory. We removed feft and right sagittal otoliths
from each individual using plastic forceps triple-washed in nitric acid (Campana et al.
2000; Brazner et al. 2004). Three independent readers aged the otolith with the most
well-defined annuli (correspondence > 90%). Prior to trace element analysis we triplerinsed otoliths in distilled, ultrapure water; air-dried them for a minimum of 24 hrs; and
stored them in acid-washed 2-mL polypropylene microcentrifuge tubes (Zeigler and
Whitledge 2010). We embedded adult otoliths in Epo-Fix epoxy and sectioned them in
the transverse plane (including the core) using a low-speed Isomet diamond saw
(Buehler, Lake Bluff, Illinois) that was cleaned with aluminum oxide lapping film after
each section to prevent contamination. Due to small size and fragilty, we placed age-0
otoliths in thermoplastic glue and ground them in the sagittal plane. We sanded all
otoliths evenly and polished them with 3M Wet or Dry sandpaper (400 grit) and
aluminum oxide lapping film, mounted them on acid-washed petrographic slides
(Donohoe and Zimmerman 2010), triple-sonicated them in ultrapure water, and dried
them in a Class 100 laminar flow hood for 24 hr.
We quantified trace element concentrations using laser ablation inductively
coupled plasma mass spectrometry (LA-ICPMS) at the Interdisciplinary Center for
Inductively Coupled Plasma Mass Spectrometry at the University of California–Davis.
86 An Agilent Technologies 7500a quadrupole ICP-MS coupled to a New Wave Research
UP-213-nm laser with He as the carrier gas was used for laser ablation analysis. Only 88Sr
and 137Ba were used to assess walleye natal origins as they were the only signatures that
were spatially variable and temporally stable (Pracheil et al. 2014). Laser parameters
were 70 % energy, 10 Hz, 40 um spot size, 25 sec dwell time, 50 sec acquisition and 25
sec background. USGS synthetic glass standard GSE-1G was used as the calibration
standard, and two additional reference standards (GSD-G1 and MACS-3) were used as
quality controls for verification of instrument accuracy and precision. Each standard was
ablated in three to five locations after every 4 samples to adjust for possible instrument
drift. Mean limits of detection for 88Sr and 137Ba were 0.01 and 0.07 ppm, respectively;
concentrations of these elements in all otoliths were well above detection limits. We
reduced raw otolith trace element data using specialized computer software (Glitter 4.4;
GEMOC CSIRO, Macquarie Research Ltd., Macquarie University, Sydney, Australia).
Otolith signatures were background subtracted, 43Ca was used as an internal standard
(Bickford and Hannigan 2005; Whitledge et al. 2007), and final trace element
concentrations were obtained by matching 43Ca counts to the sample CaO concentrations
obtained independently by electron microprobe. All otolith signatures were reported as
element:Ca ratios (μmol·mol-1) as Ca is a pseudointernal standard (Bickford and
Hannigan 2005; Ludsin et al. 2006; Whitledge et al. 2007). Only otolith signatures that
were spatially variable, temporally consistent, and positively associated with water
chemistry were used to assess walleye natal origins (Pracheil et al. 2014).
We ablated all otoliths using spot analyses. For each spot, a 15-s laser warm-up
time was followed by a 20-s dwell time during which the sample was ablated. The
87 integration time for all elements (0.01 s for 43Ca, 0.05 s for 88Sr and 137Ba) was repeated
throughout the 20-s dwell time. Following each ablation, there was a 95-sec washout
time. We quantified natal signatures of age-0 and adult walleye by ablating otoliths cores
(Ruttenberg et al. 2005). We also ablated adult otoliths at each annulus and select
interannual environments as space allowed to characterize site occupancy and movement
and entrainment patterns. This necessitated examination of recent environmental history
of age-0 walleye to construct site-specific elemental fingerprints for assessing adult
movement and entrainment (Ruttenberg et al. 2005). Water and otolith signatures were
synchronized for this analysis. When water and otolith samples were collected at the
same time, signatures were temporally matched by ablating otoliths at terminal edges,
which reflect recently occupied environments (Zeigler and Whitledge 2010, 2011).
Hereafter these signatures are referred to as “mean terminal.” When water-age-0
synchronization was not possible, age-0 individuals were invariably collected after water
samples, which permitted laser ablation at non-terminal otolith locations that
corresponded with the time of water sampling. To quantify these “adjusted mean
terminal” signatures, water and otolith chemistry were synchronized by quantifying the
number of days between water and age-0 sampling and ablating otoliths an equivalent
number of daily rings from otolith edges.
Water-otolith synchronization ensured otolith signatures represented known
capture locations and thus elemental fingerprinting was accurate for movement and
entrainment analysis. In addition, we verified that age-0 otolith signatures were consistent
at cores and edges to ensure the accuracy of our presumption that juveniles remained in
natal environments until capture. Water and otolith signatures have been matched in
88 recent studies (Zeigler and Whitledge 2010, 2011), in contrast to previous otolith
microchemistry investigations that were either unclear about temporal matching (Wells et
al. 2003, Whitledge et al. 2007) or did not involve water sampling (Brazner et al. 2004,
Cairns et al. 2004, Downs et al. 2006). Even though water chemistry need not be
measured for otolith microchemistry research, particularly in large river systems such as
the Missouri River and Eleven Point River (Bickford and Hannigan 2005) in which water
signatures are primarily driven by temporally stable geology and weathering (Gibbs
1970), we quantified water chemistry and synchronized it with age-0 otolith chemistry to
ensure reliability of our results.
Statistical Analysis
We examined water and otolith element:Ca ratios for normality and
homoscedasticity using Shapiro-Wilk tests and Levene’s tests, respectively. Water
signatures for all elements met neither assumption for parametric statistical analyses.
Thus, we compared water element:Ca ratios among sites and between years using
nonparametric Kruskal-Wallis analysis of variance by ranks, as in previous studies. We
performed post-hoc multiple comparisons using a Tukey-Kramer-Nemenyi test. We
assessed potential interactions between site and year using Friedman’s two-way analysis
of variance by ranks. Age-0 walleye terminal (i.e., mean terminal, adjusted mean
terminal) otolith signatures were normal with equal variances for some elements (e.g.,
Sr:Ca) but not others (e.g., Ba:Ca), so we used both one-way analysis of variance
(ANOVA) and Kruskal-Wallis analysis of variance by ranks to examine spatial
variability. We used least-squares linear regression (Munro et al. 2005; Zeigler and
89 Whitledge 2010; Phelps et al. 2012) to relate water and age-0 terminal otolith signatures
from each site. Statistical significance for all analyses was set at α < 0.05.
We assessed the accuracy with which bivariate (i.e., Sr:Ca, Ba:Ca) otolith
signatures classified age-0 walleye known collection site types (i.e., tributary,
embayment, mainstem) and individual sites using k-sample nearest neighbor discriminant
analysis. Used in previous microchemistry research (Bickford and Hannigan 2005), this
nonparametric method allows for reliable classification when otolith data do not meet
parametric assumptions. It assigns age-0 individuals to natal sites to which the majority
of their k nearest neighbors belong (Johnson 1998). We examined the accuracy of
different models (k = 2–8) with a leave-one-out jackknife procedure and selected the
model with the lowest error rate (k = 2) to classify adults to natal sites using the knownorigin data set (Ruttenberg et al. 2005) when Sr:Ca and Ba:Ca were in agreement, a
process that resembled a decision tree (Figure 4-2). We used juvenile walleye to develop
the discriminant model under the notion that they would reflect site-specific signatures of
capture locations unlike mobile adults. We analyzed movement data were analyzed to
identify the relative magnitude of downstream movement, upstream movement, and site
residency (i.e., neither upstream nor downstream movement) throughout all reservoirs
and within each impoundment. We summarized intra- and inter-reservoir patterns in
entrainment as percentages by site and reservoir.
Results
Water Chemistry
Water signatures varied spatially for all elemental ratios and were stable over time
for Sr:Ca, Ba:Ca, and Mg:Ca (Table 4-1, Figure 4-3). Element:Ca ratios were consistent
90 between years within sites for Sr:Ca (Friedman’s two-way analysis of variance: χ2 = 0.25,
df = 1, P = 0.62), Ba:Ca (χ2 = 1.00, df = 1, P = 0.32), and Mg:Ca (χ2 = 0.53, df = 1, P =
0.47). Sr:Ca and Mg:Ca concentrations were highest in the northernmost tributaries and
decreased moving south in accordance with longitudinal variation in surficial geology.
Non-riverine sites had comparatively lower and more consistent signatures, particularly
mainstem locations where Sr:Ca (95% CI: 4.98 ± 0.005 mmol·mol-1) and Mg:Ca (95%
CI: 787.52 ± 11.49 mmol·mol-1) were comparatively consistent. Tributary Ba:Ca
signatures increased moving downstream, but mainstem sites had lower and more
homogenous concentrations. (95% CI: 0.256 ± 0.009 mmol·mol-1).
Otolith Chemistry
Age-0 walleye terminal (i.e., mean terminal, adjusted mean terminal) otolith
signatures were variable for Sr:Ca and Ba:Ca in accordance with temporal stability
(Sr:Ca: F1,57 = 69.89, P = 0.10; Ba:Ca: F1,57 = 102.62, P = 0.49) and spatial variability
(Table 4-1) in water chemistry. Water-otolith regressions were positive and proportional
for Sr:Ca (R2 = 0.71, P < 0.01; Figure 4-4a) and Ba:Ca (R2 = 0.40, P = 0.02; Figure 44b). K-sample nearest neighbor discriminant analysis accurately classified age-0 walleye
to site types (95% mean accuracy) and individual sites (80% mean accuracy; Table 4-2).
Site-specific age-0 signatures constituted a chemical blueprint for examination of adult
environmental history. Movement
Overall, Sr:Ca and Ba:Ca resolution was sufficient to evaluate movement of adult
walleye in Missouri River reservoirs. These signatures enabled retrospective assignment
91 to natal sites, capture locations, and select environments in between. The relative
magnitudes of upstream and downstream movement were starkly divergent. Site
residency was prevalent throughout all reservoirs before the flood from 2010–2011
(48.15%, n = 78). During the flood, downstream movement (55.31%, n = 125) exceeded
upstream movement (7.96%, n = 18) and site residency (36.73%, n = 83). After the flood
from 2012–2013, downstream movement (44.98%, n = 103) and site residency (37.12%,
n = 85) were more common than upstream movement (17.90%, n = 41).
In Lake Oahe, more than 45 percent of adult walleye did not move prior to the
flood, whereas less than one third moved downstream or upstream (Table 4-3). Site
residency was most common before the flood, followed by downstream and upstream
movement. During the flood and one year later (i.e., 2012–2013), the majority of adult
walleye moved downstream or were site residents (Table 4-3).
Movement patterns of adult walleye in Lake Sharpe were somewhat different
from those in Lake Oahe. Upstream and downstream movement were relatively common
from 2010–2011 (Table 4-3). During the flood, the majority of adults were site residents
or moved downstream. Movement pattern percentages were approximately equivalent
from 2012–2013.
Adult walleye in Lake Francis Case exhibited similar movement patterns to
individuals captured in Lake Sharpe. Upstream movement and site residency were most
common before the flood, whereas site residency and downstream movement were
pervasive during the disturbance (Table 4-3). The majority of adult walleye moved
downstream after the flood from 2012–2013.
92 Movement patterns of adult walleye in Lewis & Clark Lake were distinct from
other reservoirs. A large majority of individuals were site residents before the flood,
whereas downstream movement was nearly universal during the disturbance (Table 4-3).
After the flood, adult walleye were either site residents or downstream migrants.
At a statewide scale, downstream movement to South Dakota exceeded upstream
movement to North Dakota before and during the flood. From 2010–2011, 11% (n = 18)
walleye moved downstream to South Dakota, whereas only 6% (n = 10) moved upstream
to North Dakota (Table 4-3). This disparity intensified during the flood, when 13% (n =
29) of adults moved into South Dakota and only 0.4% (n = 1) moved into North Dakota.
After the flood from 2012–2013, 3% (n = 7) moved downstream to South Dakota, and
7% (n = 16) moved upstream to North Dakota (Table 4-3).
Entrainment
Bivariate otolith element:Ca ratios enabled site assignment of age-0 and adult
walleye and comparative assessment of entrainment among reservoirs and sites. Adults
collected in Lake Sharpe had natal otolith signatures characteristic of Lake Oahe
tributaries (Figure 4-5a, 4-5b), indicating entrainment through Oahe Dam. Similarly,
adults collected in Lake Francis Case had natal otolith signatures from Lake Sharpe sites
(Figure 4-6a, 4-6b), confirming entrainment through Big Bend Dam. Entrainment
increased moving downstream such that adults collected in Lewis & Clark Lake had natal
otolith signatures characteristic of sites from lakes Oahe, Sharpe, and Francis Case
(Figure 4-7a, 4-7b).
A summary of adult walleye entrainment by reservoir and age class provided
93 powerful insight into flood effects germane for management. One-fourth (25.19%, n =
34) of adult walleye collected in lakes Sharpe, Francis Case, and Lewis & Clark were
entrained at some time in their lives. Twelve of these individuals were entrained multiple
times, resulting in 48 total entrainment events. The frequency of entrainment increased
moving downstream in Missouri River reservoirs (Table 4-4). One-fifth (18.87%, n = 10)
of adult walleye from Lake Sharpe were entrained, compared to one-fourth (24.39%, n =
10) from Lake Francis Case and one-third (34.15%, n = 14) from Lewis & Clark Lake.
The 2011 flood caused extensive entrainment of adult walleye. Nearly two-thirds
(64.71%; n = 22) of all entrained individuals moved through dams during the disturbance,
accounting for 45.83 percent of all entrainment events. Entrainment occurred much less
frequently in other annual periods, the highest of which were 2012–2013 (n = 8 events,
16.67%) and and 2007–2008 and 2010–2011 (n = 5 events, 10.42%). Flood-induced
entrainment increased moving downstream (Table 4-4). Locations with the highest
entrainment percentages by reservoir included West Bend in Lake Sharpe (100.00%, n =
3), Chamberlain Area in Lake Francis Case (29.41%, n = 5), and Gavin’s Point Dam in
Lewis & Clark Lake (42.86%, n = 6; Table 4-5). High percentages of flood-entrained
adult walleye moved into and were collected in Lake Sharpe (57.14%, n = 4) and Lake
Francis Case (60.00%, n = 6) during the flood, whereas only 29.00 % (n = 4) entered
Lewis & Clark Lake. The majority (68.18%, n = 15) of entrained individuals were age3+, but low capture efficiency for age-2 walleye may have contributed to a low
entrainment estimate for this age class (31.82%, n = 7).
94 Discussion
Movement has important implications for research and management of fisheries
throughout the world (Lucas and Baras 2000, Pelicice and Agostinho 2008, Baumgartner
et al. 2014). Knowledge of movement is critical for identifying and protecting nursery
areas (Landsman et al. 2011), evaluating and improving stocking programs (Fielder
2002), and controlling invasive species (Christie and Goddard 2003). Walleye movement
research is extensive but has generally focused on limited topics (e.g., reproductive
biology, stocking, and habitat use; Crowe 1962) in lacustrine systems (e.g., Great Lakes;
Wang et al. 2007) using mark-recapture and radiotelemetry. Emerging technologies offer
new avenues for investigating walleye movement and other aspects of environmental
history (e.g., entrainment, natal origins). Otolith microchemistry is one such method that
has facilitated environmental history research on fish natal origins (Zeigler and
Whitledge 2010, 2011) and movement (Allen et al. 2009). Other topics have included
stock composition (Bickford and Hannigan 2005), habitat fingerprinting (Brazner et al.
2004), and invasion biology of nonnative species (Munro et al. 2005). However, otolith
microchemistry has not been used to examine environmental history and flood response
of walleye, a knowledge gap filled by this study.
Our results add to a growing body of literature demonstrating the efficacy of
otolith microchemistry for fisheries management throughout the world (Campana et al.
2000, Ruttenberg et al. 2005, Rohtla et al. 2014). Our findings indicated otolith
microchemistry is a reliable tool for examining walleye movement and entrainment in
Missouri River reservoirs, corroborating previous studies on diverse species (e.g.,
American eel [Anguilla rostrate], Cairns et al. 2004; beaked redfish [Sebastes mentella
95 and S. fasciatus], Campana et al. 2007; green sturgeon [Acipenser medirostris], Allen et
al. 2009). Spatial variability and temporal stability in water Sr:Ca and Ba:Ca signatures
and positive, proportional water-otolith relationships laid a foundation for microchemical
analysis (Zeigler and Whitledge 2010, 2011; Gahagan et al. 2012). Mean terminal otolith
Sr:Ca and Ba:Ca ratios of age-0 walleye were spatially heterogeneous in accordance with
water chemistry, representing elemental fingerprints for adult site assignment (Gahagan
et al. 2012). However, Mg:Ca, Mn:Ca, and Na:Ca did not reflect water chemistry and
were thus unreliable signatures. Mg and Na are susceptible to chemical alteration as they
are loosely bound in the CaCO3 matrix compared to Sr and Ba (Milton and Chenery
1998, Proctor and Thresher 1998, Hedges et al. 2004), whereas Mn concentrations are
typically low (Wells et al. 2003) and unstable (Wetzel 1983) due complex metabolic
processes (Gibson-Reinemer et al. 2009) and growth-related biogeochemical changes
(Limburg et al. 2010). Overall, we anticipate Sr:Ca and Ba:Ca to be most effective for
future otolith microchemistry studies in Missouri River reservoirs. Our results fill a gap
in the scientific literature as otolith microchemistry has scarcely been used to investigate
fish entrainment (Sandin et al. 2005; Shima and Swearer 2009) or walleye environmental
history (Bickford and Hannigan 2005).
Walleye movement has traditionally been studied using mark-recapture and
radiotelemetry (Landsman et al. 2011), whereas emerging methods such as otolith
microchemistry have been used scarcely (Bickford and Hannigan 2005). Herein, we
demonstrated that trace element signatures of Missouri River tributaries are distinct and
represent a powerful tool for discerning intra- and inter-reservoir movements of walleye.
Site classification using k-sample nearest neighbor discriminant analysis was highly
96 accurate for tributaries, particularly those in Lake Oahe, reflecting bivariate distinctness
in water chemistry resulting from heterogeneous surficial and bedrock geology.
Classification was also accurate for embayments and mainstem locations, indicating
Sr:Ca and Ba:Ca are reliable signatures for retrospective assessment of environmental
history. Overall, otolith microchemistry is an auspicious alternative to traditional markrecapture and radiotelemetry studies for examining walleye movement and entrainment in
Missouri River reservoirs.
Walleye movement within and among impoundments was dominated by
downstream passage and site residency, whereas upstream movement was relatively
uncommon. The pervasiveness of downstream movement likely resulted from high
discharges (Spoelstra et al. 2008) but may have also reflected the northerly orientation of
critical recruitment areas, particularly tributaries, within reservoirs. Site residency of
percids has been documented previously (Spoelstra et al. 2008) and may indicate
adequate prey resource availability and/or spawning site fidelity during stable
hydrological regimes (i.e., before and after floods). Walleye are known to move upstream
before (Paragamian 1989) and during (DePhilip et al. 2005) spawning season. Despite the
prevalence of tributary spawning observed in this study, upstream spawning movements
are evidently less important than downstream movements and site residency in Missouri
River reservoirs. However, notable exceptions existed, as Sr:Ca and Ba:Ca signatures
indicated some individuals moved upstream from the Moreau River in South Dakota to
the Cannonball River in North Dakota prior to flood-induced downstream movement.
The temporal scale of our assessment (i.e., years) may also have masked intra-annual
upstream movement. Finer-scale analyses would enhance future otolith microchemistry
97 research in Missouri River reservoirs.
Although downstream movement of walleye was pervasive during the flood,
movement patterns were variable among reservoirs before and after the disturbance, as
has been documented in previous research in the Great Lakes (Wang et al. 2007). Site
residency was common before the flood in Lake Oahe, whereas downstream movement
was prevalent after the disturbance. Site residency may have reflected the high abundance
of tributaries (e.g., spawning sites), tributary site fidelity, and/or abundant prey resources.
Compared to Lake Oahe, upstream movement was more common in lakes Sharpe and
Francis Case before the flood, which may have reflected the northerly orientation of
spawning sites (i.e., Bad River in Sharpe, White River and Chamberlain Area in Francis
Case) in these reservoirs. Walleye exhibited pre-flood and post-flood site residency in
Lewis & Clark Lake, likely indicating the importance of the Lewis & Clark Delta as a
spawning and nursery area (Kaemingk et al. 2007, Graeb et al. 2009). On a statewide
scale, downstream movement from North Dakota to South Dakota was more extensive
than upstream movement from South Dakota to North Dakota. Because North Dakota
and South Dakota natal sites contribute walleye to the opposite state and individuals
move extensively between states, collaborative interstate management is important. We
recommend that managers ensure that harvest regulations (i.e., creel limits, slot limits,
one-over rules) are compatible among states under normal hydrological conditions and
after floods. For example, because floods impede upstream movement and cause
downstream movement and uneven spatial distribution of walleye, post-flood regulations
should be relatively conservative and liberal in North Dakota and South Dakota,
respectively. In addition, we advise managers to collaborate on habitat protection and
98 restoration activities (e.g., riparian zone preservation/enhancement, sediment dredging),
particularly at important spawning locations (e.g., Moreau and Cannonball rivers).
Moreover, we recommend managers in both states jointly conduct human dimensions
research to understand stakeholders in both states and use this information to design
riverscape (Fausch et al. 2002) management strategies and public engagement initiatives.
Temporal patterns in entrainment revealed herein may also advance walleye
management in Missouri River impoundments. An important observation is that
entrainment is not confined to floods. Baseline entrainment suggests walleye populations
in Missouri River reservoirs are more connected than previously believed and indicates
the importance of riverscape management. For example, large-scale habitat protection
and restoration programs are likely to enhance walleye populations inside and outside the
specific reservoirs where they are implemented. Entrainment increased during the flood
and moving downstream in Missouri River reservoirs. This indicates walleye survive dam
passage and entrainment increases under high discharge, as with saugeye (female walleye
X male sauger [Sander canadensis]; Spoelstra et al. 2008) and other species (e.g., striped
bass [Morone saxatilis], delta smelt [Hypomesus transpacificus]; Grimaldo et al. 2009).
In addition, otolith microchemistry represents a predictive tool to estimate entrainment
during floods and normal hydrological regimes. We recommend that managers enact
harvest regulations that are conservative in upper reservoirs (i.e., Oahe, Sharpe)
compared to lower reservoirs, especially after floods in the vicinity of important natal
sites. Although less conservative regulations in lower reservoirs are likely justifiable after
floods due to extensive downstream movement, managers should monitor entrainment
from these impoundments (e.g., from Lake Lewis & Clark to the Missouri River outside
99 South Dakota) to ensure populations are not vulnerable to liberal harvest. However, if
entrainment subsidization increases abundance in lower reservoirs such that population
dynamics show density-dependent responses (e.g., decreased growth, increased
mortality), we recommend managers liberalize regulations to decrease density. Much like
baseline entrainment, the fact that lower reservoirs, with fewer important natal sites,
contain higher proportions of entrained walleye signifies the significance of habitat
conservation in upper reservoirs where natal environments are more abundant.
Given that managers can choose to mitigate entrainment (e.g., strobe lights, sound
deterrents), we project potential outcomes and implications under different management
scenarios. Currently in the absence of entrainment mitigation, the Missouri River
functions as a riverscape of connected impoundments with interconnected walleye
populations. Entrainment is known to occur under all hydrological conditions, and it
increases with floods and moving downstream. As such, reservoirs essentially function as
modified put-grow-take systems that are replenished by upstream sources. In general, this
situation calls for liberal walleye management, but we recommend that decision makers
monitor entrainment to confirm it continually restocks populations. In addition, we advise
managers to quantify site-specific spawning-recruitment relationships and protect
upstream populations via habitat conservation to ensure they can replenish lower ones.
Moreover, when certain length categories are more vulnerable to entrainment than others,
we suggest managers design regulations that promote harvest of these size classes, but
not to the extent of eliminating entrainment and the downstream subsidies it provides.
In contrast, if managers decided to mitigate entrainment, they would rely on
natural reproduction to replenish populations. Reservoirs with abundant natal habitats
100 (i.e., Oahe) may be relatively self-sustaining, but lower reservoirs with high proportions
of entrained individuals and fewer natal environments may not be. Our results suggest
that entrainment plays a vital role in structuring walleye populations in Missouri River
reservoirs. For example, 47.5% of adult walleye in Lake Francis Case hatched in
upstream reservoirs, as did 35% of adults in Lake Lewis & Clark. Although these
reservoirs contain natal sites, our results suggest natural reproduction may not match
entrainment subsidization. We recommend managers compare the advantages and
disadvantages of extant entrainment in relation to goals, which will promote sciencebased walleye management in Missouri River reservoirs.
Overall, otolith microchemistry is an informative, high-resolution tool for
elucidating walleye movement and entrainment in Missouri River reservoirs. The
technique provides reliable information on intra- and inter-reservoir movements that may
be used by agencies to advance walleye management. We recommend future researchers
examine entrainment susceptibility of specific walleye age classes, particularly age-1 and
age-2, which were difficult to collect in the present study likely due to impaired spawning
and a poor 2011 year class during the flood (Gebken and Wright 1972; Serns 1982). In
addition, we advise future researchers to evaluate natal origins, movement, and
entrainment of other species at statewide and finer scales. Additional otolith signatures,
including trace elements (e.g., 24Mg) and/or isotopes (e.g., δ13C, δ18O, 87Sr/86Sr), may be
useful for these analyses. Otolith microchemistry may also inform stock discrimination
studies (Campana et al. 2000; Bickford and Hannigan 2005), especially those focused on
invasive species. The technique could provide vital provenance information to streamline
control and eradication efforts as invasive fishes threaten ecosystems and economies.
101 Finally, otolith microchemistry may advance conservation programs for threatened and
endangered fishes (e.g., pallid sturgeon [Scaphirhychus albus]) by furnishing information
on natal origins, movement, and entrainment.
Acknowledgments
We thank R. Johnston, C. Hayer, T. Rapp, and J. Mecham of South Dakota State
University for field assistance. We thank B. Hanten, H. Meyer, K. Potter, G. Knecht, J.
Sorensen, T. Sorensen, R. Trible, and B. Larson of the South Dakota Department of
Game, Fish and Parks and P. Bailey of the North Dakota Game and Fish Department for
technical advice and assistance. We thank A. Shiller and lab members at the University of
Southern Mississippi for providing water sampling kits and measuring trace element
concentrations in water samples. We thank G. Barford, J. Commisso, and J. Glessner at
the University of California–Davis for assistance with otolith microchemistry analyses.
Funding for this project was provided by the Federal Aid in Sport Fish Restoration
program, Project 3M3328, Study 1526, administered by the South Dakota Department of
Game, Fish and Parks, United States Fish & Wildlife Service, South Dakota State
University Department of Natural Resource Management, and South Dakota Agricultural
Experiment Station.
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110 Tables
Table 4-1. Results of one-way ANOVA testing for spatial variability in water
element:Ca ratios among Missouri River tributaries and sites within reservoirs. A
significant P value (≤ 0.05) indicates water chemistry is spatially variable in the
associated water body.
Element Water body
Sr:Ca
Tributaries
Ba:Ca
Mg:Ca
Mn:Ca
Na:Ca
F
F8,6 = 2155.50
P
< 0.01
Oahe tributaries
F5,3 = 73.90
< 0.01
Oahe
F15,13 = 1914.70 < 0.01
Sharpe
F7,6 = 169.28
< 0.01
Francis case
F7,8 = 316.02
< 0.01
Lewis & clark
F6,6 = 1528.10
< 0.01
Tributaries
F8,6 = 7468.80
< 0.01
Oahe tributaries
F5,3 = 1750.80
< 0.01
Oahe
F15,13 = 747.79
< 0.01
Sharpe
F7,6 = 3087.40
< 0.01
Francis case
F7,8 = 158.64
< 0.01
Lewis & clark
F6,6 = 817.86
< 0.01
Tributaries
F8,6 = 5093.40
< 0.01
Oahe tributaries
F5,3 = 1017.20
< 0.01
Oahe
F15,13 = 792.22
< 0.01
Sharpe
F7,6 = 16.68
< 0.01
Francis case
F7,8 = 4529.30
< 0.01
Lewis & clark
F6,6 = 459.41
< 0.01
Tributaries
F8,6 = 32.95
< 0.01
Oahe tributaries
F5,3 = 4723.10
< 0.01
Oahe
F15,13 = 349.26
< 0.01
Sharpe
F7,6 = 703.18
< 0.01
Francis case
F7,8 = 313.41
< 0.01
Lewis & clark
F6,6 = 493.06
< 0.01
Tributaries
F8,6 = 6803.30
< 0.01
Oahe tributaries
F5,3 = 2576.30
< 0.01
Oahe
F15,13 = 2132.50 < 0.01
Sharpe
F7,6 = 142.93
< 0.01
Francis case
F7,8 = 1080.90
< 0.01
Lewis & clark
F6,6 = 1086.77
< 0.01
111 Table 4-2. Results of k-sample nearest neighbor discriminant analysis with leave-one-out
jackknife cross-validation using Sr:Ca and Ba:Ca ratios of age-0 walleye. Data include
the number known to have come from site types and individual sites (Known), the
number assigned to those locations (Assigned), the percent deviation of Assigned from
Known (Deviation), and the percentage of individuals classified to known locations
(Accuracy). Signatures were measured at otolith locations synchronized with water
sample collection to ensure reliable spatial blueprinting.
Natal origin
Site type
Tributary
Embayment
Mainstem
Site
Bad R.
Beaver Bay
Cannonball R.
Cheyenne R.
Ft. George
Heart R.
Knife R.
Minneconjou
Moreau R.
N. Bay
Platte Creek
Springfield
W. Pollock
W. Whitlocks
Known (number)
Assigned (number)
Deviation (%)
Accuracy (% of known)
16
28
2
19
26
3
19
-7
50
100
86
100
Mean: 95
4
2
2
4
2
2
2
4
2
8
4
2
3
5
7
2
2
3
3
3
2
4
2
8
3
3
1
3
75
0
0
-25
50
50
0
0
0
0
-25
50
-67
-40
100
100
100
75
100
100
100
75
100
63
50
100
33
20
Mean: 80
.
112 Table 4-3. Percent (number) of adult walleye exhibiting downstream movement,
upstream movement, and site residency at reservoir (Oahe, Sharpe, Francis Case, Lewis
& Clark) and statewide (North Dakota [ND], South Dakota [SD]) spatial scales. At the
statewide scale, “downstream” and “upstream” denote movements to SD and ND,
respectively.
Year
Reservoir/state
2010–2011 Oahe
Sharpe
Francis case
Lewis & clark
ND
SD
2011–2012 Oahe
Sharpe
Francis case
Lewis & clark
ND
SD
2012–2013 Oahe
Sharpe
Francis case
Lewis & clark
ND
SD
Downstream
32 (24)
42 (13)
27 (6)
6 (2)
-11 (18)
53 (50)
40 (21)
41 (16)
93 (38)
-13 (29)
53 (51)
31 (16)
58 (23)
32 (13)
-3 (7)
Upstream
21 (16)
48 (15)
32 (7)
3 (1)
6 (10)
-2 (2)
17 (9)
15 (6)
2 (1)
< 1 (1)
-19 (18)
33 (17)
15 (6)
0 (0)
7 (16)
--
Residency
47 (36)
10 (3)
41 (9)
91 (30)
94 (152)
89 (144)
45 (42)
42 (22)
44 (17)
5 (2)
99 (225)
87 (197)
28 (27)
36 (19)
27 (11)
68 (28)
93 (213)
97 (222)
113 Table 4-4. Summary of adult walleye entrainment in Missouri River reservoirs, where %
Flood-entrained represents the percentage of all fish and entrained fish (in parentheses)
that were entrained during the 2011 flood. Similarly, % Baseline represents the
percentage of all fish and entrained fish that were not entrained during the 2011 flood.
Reservoir
# Ablated
Sharpe
53
Francis case
41
Lewis & clark
41
Total
135
# Entrained
10
10
14
34
% Entrained % Flood-entrained
18.87
13.21 (70.00)
24.39
17.07 (70.00)
34.15
19.51 (57.14)
25.19
16.30 (64.71)
% Baseline
5.66 (30.00)
7.32 (30.00)
14.63 (42.86)
8.88 (35.29)
114 Table 4-5. Summary of adult walleye entrainment by collection site in Missouri River
reservoirs. Entrained (flood) represents all entrained and 2011 flood-entrained individuals
(in parentheses).
Reservoir
Sharpe
Sharpe
Sharpe
Sharpe
Francis case
Francis case
Francis case
Lewis & clark
Lewis & clark
Lewis & clark
Site
# ablated # entrained (flood) % entrained (flood)
Fort george
17
1 (1)
5.88 (100.00)
N. shore
16
2 (1)
12.50 (50.00)
Stilling basin
17
4 (2)
23.53 (50.00)
W. bend
3
3 (3)
100.00 (100.00)
Stilling basin
17
5 (4)
29.41 (80.00)
N. bay
13
2 (1)
15.38 (50.00)
Platte creek
11
3 (2)
27.27 (66.67)
Gavins point dam
14
6 (4)
42.86 (66.67)
Tailrace
11
3 (2)
27.27 (66.67)
LC delta
16
5 (2)
31.25 (40.00)
115 Figures
Figure 4-1. Water and walleye sampling locations in Missouri River reservoirs. All sites
were sampled for water trace element concentrations using a syringe-filtration protocol
described in Shiller (2003). Shapes denote locations where fish of different ages were
sampled: right-angle triangles (age-0), upright triangles (adult), diamonds (both).
116 Figure 4-2. A conceptual model for evaluating movement and entrainment of walleye in
Missouri River reservoirs. Site assignment of adult walleye to locations in the Missouri
River resembled a decision tree involving synthesis of bivariate water and otolith
signatures, the river-wide water-otolith relationship, collection locations, and reservoir
orientation. The water-otolith relationship generated from mean terminal otolith
signatures of age-0 walleye collected in fall 2012.
117 Figure 4-3. Mean water (a) Sr:Ca and (b) Ba:Ca value at the 14 sites where both water
and age-0 walleye were collected in late summer 2012. Means with the same letter are
not significantly different (ANOVA with Tukey’s HSD test on log10 transformed values;
P ≤ 0.05). Error bars represent SEs.
118 Figure 4-4. Linear regression of age-0 walleye mean terminal otolith (a) Sr:Ca and (b)
Ba:Ca signatures on water ratios at collection sites lakes Oahe, Sharpe, Francis Case, and
Lewis & Clark. Fish and water sampling occurred in late summer 2012. Error bars
represent 1 SEM.
119 Figure 4-5. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye
collected in Lake Sharpe, South Dakota, USA in Summer 2013. Site assignments reflect
bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black circle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others
(e.g., white triangle) exhibited variable signatures suggesting intra-reservoir movement
and entrainment through mainstem dams. All transects proceed from the core of the
otolith toward the edge. Individuals depicted here were collected at Sharpe Stilling Basin.
120 Figure 4-6. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye
collected in Lake Francis Case, South Dakota, USA in Summer 2013. Site assignments
reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black triangle)
displayed relatively stable element:Ca ratios indicating prolonged site residence, whereas
others (e.g., white triangle) exhibited variable signatures suggesting intra-reservoir
movement and entrainment through mainstem dams. All transects proceed from the core
of the otolith toward the edge. Individuals depicted here were collected near
Chamberlain, South Dakota.
121 Figure 4-7. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye in
Lewis & Clark Lake, South Dakota, USA in Summer 2013. Site assignments reflect
bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., white circle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others
(e.g., black circle) exhibited variable signatures suggesting intra-reservoir movement and
entrainment through mainstem dams. All transects proceed from the core of the otolith
toward the edge. Individuals depicted here were collected at Gavins Point Dam.
122 CHAPTER 5: EFFECTS OF CATASTROPHIC FLOODING ON A DELTA
ECOSYSTEM IN THE MISSOURI RIVER
This chapter is being prepared for submission to Ecology of Freshwater Fish and was coauthored by Mark J. Fincel, Christopher M. Longhenry, and Brian D. S. Graeb.
123 Abstract
Catastrophic flooding in the Missouri River in 2011 provided an opportunity to
measure immediate ecological effects of high discharge in the Lewis and Clark Delta,
South Dakota. We evaluated trends in structural indices and relative abundance of the
Delta fishes semiannually from 2004 to 2012 and identified effects of flooding on the fish
community and aquatic habitats. Community evenness (J’) consistently decreased from
0.882 to 0.725 before the flood but rose to 0.835 after the disturbance. Pre-flood species
richness declined from 25 to 15, whereas post-flood richness stabilized above 20.
Diversity (Fisher’s α) decreased from 4.56 to 3.48 before the flood and stabilized at 3.27
after the disturbance, indicating absence of an immediate structural flood response.
Nonmetric multidimensional scaling provided insight into flood effects on relative
abundance of specific community segments. A majority of species were more abundant
after the flood regardless of native/nonnative status, trophic guild, and length. Relative
abundance of juvenile freshwater drum Aplodinotus grunniens and white crappie
Pomoxis annularis declined from pre-flood levels, whereas catch-per-unit-effort of smallbodied native species remained stable. Channel and backwater frequency and main
channel width in the Delta declined after the flood, whereas sandbars became more
abundant. Habitat changes likely had minimal effects on the fish community as
availability of shallow-water habitats was still higher in the Delta compared to main
channels depauperate in slackwater areas necessary for spawning. Our results suggest the
Delta serves as a refuge environment for juvenile and adult fishes during and
immediately after catastrophic flooding.
Key words: flooding, Missouri River, Lewis & Clark Delta, fish, habitat
124 Introduction
Floods cause environmental variability and structure biotic communities in large
floodplain rivers (Poff et al., 1997; Ward et al., 1999), yet long-term research examining
temporal effects of disturbance is scarce (Bischoff and Wolter, 2001). Floods connect
rivers with floodplains and supply nutrients and organic matter to aquatic-terrestrial
transition zones rich in biotic diversity (Ward and Stanford, 1995). Lateral riverfloodplain connectivity generates an edge effect whereby a dynamic littoral zone
characterized by diversity in flow and temperature stimulates biological productivity and
habitat heterogeneity (Junk et al., 1989; Tockner et al., 2000). Connectivity is especially
important in freshwater deltas, which are among the most biologically productive and
ecologically diverse ecosystems and depend on sediment-laden river water to maintain
their habitat complexity and biodiversity (Long and Pavelsky, 2013). Frequent flooding
in the Missouri River diminished after impoundment during the mid-twentieth century,
depleting sediment inputs to deltas. However, disturbances that simulate historic floods
and reestablish river-floodplain connectivity still occur, providing opportunity to evaluate
physical and biological effects in delta ecosystems.
Effects of floods on aquatic ecosystems are mediated by diverse factors including
magnitude, timing, and frequency of disturbance. These factors determine flood strength
and direction of abiotic and biotic responses (Kano et al., 2011). Population- and
community-level effects are driven by abiotic processes when discharge fluctuates,
whereas biotic processes such as competition and predation dominate under stable flow
regimes (Poff and Ward, 1989). Responses of aquatic organisms to floods are also
governed by degree of environmental adaptation (Bischoff and Wolter, 2001). Over
125 evolutionary time scales, morphological (e.g., dorsoventral orientation), physiological
(e.g., turbidity tolerance, swimming strength), and behavioral (e.g., habitat preference)
adaptations enable population recovery, stability, and persistence after extreme
disturbance (DeBoer et al., 2011; Kano et al., 2011; Valdez et al., 2011).
An historic flood occurred in the Missouri River in 2011. Late-melting snow in
the Rocky Mountains and late-spring rainfall in the Great Plains caused catastrophic
flooding in spring and summer. Runoff totaled 75.2 billion m3 and exceeded system
capacity by > 20 percent above Sioux City, IA (USACE, 2012). All six mainstem
reservoir dams recorded record discharge. In addition to social, economic, and
hydrological effects, the disturbance may have caused ecological changes. Preliminary
research indicates substantial physical alterations, including bed degradation, thalweg
migration, and loss of river-floodplain connectivity (Cowman, 2012), but biotic effects
remain largely unknown.
Freshwater deltas, formed by prolonged sedimentation at tributary mouths and
upper reaches of reservoirs (Johnson, 2002), are ideal environments for investigating
biotic effects of floods. Deltas in the Missouri River contain warm, turbid water and
heterogeneous habitats that simulate the historic riverscape (Graeb et al., 2009) and
support diverse fish communities (Kaemingk et al., 2007). Despite their ecological
functions, deltas decrease reservoir storage capacity and impede waterfront access. The
USACE is examining feasibility of sediment management alternatives (i.e., dewatering
and sediment flushing, channelization, dredging, tributary diversion) to mitigate
sedimentation in deltas (USACE, 2011). Our objective was to treat the 2011 flood as a
natural experiment in the Lewis and Clark Delta to evaluate ecological effects of
126 discharges used in potential management scenarios (e.g., dewatering and sediment
flushing). We evaluated changes in structural indices and relative abundance of the Delta
fish community and alterations in aquatic habitats resulting from the flood. This
information may promote science-based aquatic resource management in the Delta in the
future.
Methods
Study Area
Lewis and Clark Lake is the smallest and southernmost mainstem Missouri River
impoundment (Figure 5-1). Formed after completion of Gavins Point Dam in 1955, it has
a surface area of 105 km2 (USACE, 2006), maximum depth of 16.7 m, and mean depth of
5.0 m (Wickstrom, 2004). Approximately 70 km of riverine habitat extend upstream from
the reservoir to Fort Randall Dam, which impounds Lake Francis Case.
The Lewis and Clark Delta (hereafter Delta) extends 34 km above Lewis and
Clark Reservoir, and the Missouri National Recreation River, a remnant reach,
encompasses the remaining 36 km to Fort Randall Dam (Graeb et al., 2009). The Delta is
a complex ecosystem characterized by diverse habitats, warm temperatures, high
turbidity, and connectivity to the floodplain (Graeb et al., 2009). It was created by
sediment deposition from the Niobrara River, which annually delivers 2.57 million metric
tons of sediment to Lake Lewis and Clark (USACE, 2001), causing the Delta to expand
by 105 m (Elliott and Jacobson, 2006) and lose 2,400 acre-feet of water storage per year
(USACE, 2011). Gauge height in the Delta during the 2011 flood was the highest on
record since 1980 (Figure 5-2).
127 Fish Sampling
The Delta fish community was sampled for nine consecutive years from 2004 to
2012. Juvenile fishes were collected in shallow environments with modified mini-fyke
nets (MF; 1.2 m by 0.6 m frames, 3.2 mm bar mesh; Schreck 2010) two or three times per
summer, including the flood in 2011. Adult fishes and small-bodied native species were
sampled in nearshore habitats with boat-mounted electrofishing (EF) (Coffelt VVP-15
control unit; C-phase, pulsed-DC current) in spring, summer, and autumn of 2005, 2008,
2009, and 2012.
Aquatic Habitats
Changes in aquatic habitats in the Delta resulting from the 2011 flood were
evaluated using Landsat landcover geodatabases provided by the USACE. Pre-flood
(June 2010) and post-flood (October 2011) imagery was displayed into ARCMap 10.1
and a point shapefile containing 30 equidistantly-spaced transect lines (perpendicular to
streamflow; Appendix K) between the Niobrara River confluence and downstream edge
of the Delta was overlaid onto each of the images. Channels, backwaters, and sandbars
were enumerated and main channel width and largest-backwater area were measured
along each transect.
Statistical Analysis
Pre-flood EF catch-per-unit-effort (CPUE) data from 2005 (Kaemingk et al.,
2007) and 2008-2009 (Schreck, 2010) were combined with post-flood data from 2012.
Pre-flood, flood, and post-flood MF data spanning from 2004-2012 were compiled from
128 the United States Fish and Wildlife Service (D. James, USFWS, unpublished data).
Fishes were categorized into seven groups based on length category (i.e., juvenile, adult;
Gabelhouse, 1984) and introduction history to assess temporal community dynamics:
juvenile sport, juvenile native, juvenile introduced, adult sport, adult native, adult
introduced, and small-bodied native (Schreck, 2010). Fishes were also categorized into
groups based on trophic guild (e.g., detritivore, invertivore, omnivore, piscivore; Table 51).
Delta fish community and aquatic habitat data were analyzed using univariate and
multivariate statistical methods. Temporal trends in structural indices (e.g., species
richness, evenness [J’], and diversity [Fisher’s α]) were assessed in 2005, 2008, 2009,
and 2012 with years assigned to flood periods (i.e., pre-flood, flood, post-flood). The
entire species pool was condensed to common species (i.e., > 1 percent of total catch at
sampling sites) to remove effects of rare species and species difficult to collect. Because
sample sizes differed among assemblages, rarefaction was performed to compare species
richness over time (Kwak and Peterson, 2007). Temporal trends in structural indices were
evaluated using a bootstrapping procedure in program PAST (Paleontological Statistics)
analogous to a two-sample t-test, which was used to compare pre-flood and post-flood
frequencies of habitat types (e.g., channels, backwaters, sandbars), mean main channel
width, and mean largest-backwater area along georeferenced transects throughout the
Delta.
Multivariate statistical analyses were performed to assess temporal changes in
relative abundance of the Delta fish community. All fishes other than small-bodied native
species were categorized by length, introduction history, and trophic guild to evaluate
129 effects of the flood on different community segments. Non-metric multidimensional
scaling (NMS) using square-root-transformed Bray-Curtis distance and post-hoc
permutational multivariate analysis of variance (PERMANOVA, N = 1000 permutations)
was performed in PRIMER (PRIMER-E Ltd, Plymouth, United Kingdom; Clarke and
Gorley, 2006) to assess multivariate trends in relative abundance among flood periods for
length, introduction history, and trophic guild groupings. Significance was set at α = 0.05.
One-way analysis of variance (ANOVA) with post-hoc Tukey’s Honestly Significant
Difference (HSD) tests were used to evaluate effects of flooding on relative abundance of
individual species by age category. Mini-fyke net data was only used for juvenile
assessments, whereas EF data were used for analyses of adults and small-bodied native
species due to higher catch rates (Table 5-2).
Results
Structural Indices
More than seven thousand fishes (N = 7,280) representing 37 species were
collected in the Lewis and Clark Delta from 2004-2012. Rarefied species richness
decreased from 2005 (25) to 2008 (15) but was unaffected by the flood, remaining stable
from 2009 (22) to 2012 (21; Table 5-3). Evenness (J’) declined from 2005 (0.882) to
2008 (0.796) to 2009 (0.725) but rose in 2012 (0.835) after the flood (Table 5-3).
Diversity (Fisher’s α) decreased from 2005 (4.56) to 2008 (2.17) but stabilized thereafter
(3.48 in 2009, 3.27 in 2012; Table 5-3), indicating absence of an immediate structural
flood response.
130 Relative Abundance by Introduction History
Relative abundance of juvenile and adult introduced fishes varied among flood
periods, driven by increased post-flood CPUE of Common Carp (Cyprinus carpio)
juveniles (F = 20.22, df = 2, 69, p < 0.01) adults (F = 18.22, df = 1, 174, p < 0.01).
Juvenile native species exhibited distinct relative abundance among flood periods as
CPUE of Freshwater Drum (Aplodinotus grunniens) declined (F = 4.43, df = 2, 69, p <
0.01) and CPUE of River Carpsucker (Carpiodes carpio) increased (F = 50.78, df = 2,
69, p < 0.01; Figure 5-3). Relative abundance of adult native species varied among flood
periods due to increased CPUE of Freshwater Drum (F = 59.59, df = 1, 174, p < 0.01),
River Carpsucker (F = 17.40, df = 1, 174, p < 0.01), and Shorthead Redhorse
(Moxostoma macrolepidotum; F = 15.66, df = 1, 174, p < 0.01; Figure 5-4).
Relative abundance of juvenile sport species differed among flood periods as
CPUE increased for Bluegill (Lepomis macrochirus; F = 12.53, df = 2, 69, p < 0.01) and
Black Crappie (Pomoxis nigromaculatus; F = 19.70, df = 2, 69, p < 0.01) and decreased
for White Crappie (Pomoxis annularis; F = 6.08, df = 2, 69, p < 0.01) and Smallmouth
Bass (Micropterus dolomieu; F = 5.48, df = 2, 69, p < 0.01; Figure 5-3). Similarly, CPUE
of adult sport species was heterogeneous among flood periods, driven by increases in
relative abundance of Bluegill (F = 7.00, df = 1, 174, p < 0.01), Channel Catfish
(Ictalurus punctatus; F = 11.37, df = 1, 174, p < 0.01), Largemouth Bass (Micropterus
salmoides; F = 17.36, df = 1, 174, p < 0.01), and Sauger (Sander canadensis; F = 5.58, df
= 1, 174, p = 0.02; Figure 5-4). Small-bodied native species displayed stable relative
abundance across flood periods as CPUE remained consistent for Emerald Shiner
(Notropis atherinoides; F = 1.57, df = 2, 69, p = 0.22), Red Shiner (Cyprinella lutrensis;
131 F = 0.05, df = 2, 69, p = 0.82), and Spotfin Shiner (Cyprinella spiloptera; F = 1.98, df =
2, 69, p = 0.16; Figure 5-4).
Relative Abundance by Trophic Guild
Juvenile detritivores exhibited distinct relative abundance among flood periods,
caused by increased post-flood CPUE of River Carpsucker (Figure 5-3). Adult
detritivores exhibited a similar trend as inconsistency in CPUE among flood periods (F =
3.88, df = 63, p < 0.01) was attributable to River Carpsucker (Figure 5-4).
Relative abundance of invertivores was heterogeneous among flood periods (F =
2.86, df = 2, 169, p = 0.03) and within seasons (F = 2.47, df = 2, 169, p = 0.05) despite
stability for SBN species, which were invertivorous in the Delta. Variability was caused
by adult Shorthead Redhorse, which were more abundant after the flood than before
(Figure 5-4).
Juvenile omnivores exhibited uneven relative abundance among flood periods as
post-flood CPUE was higher for Common Carp, Bluegill, and Black Crappie and lower
for Freshwater Drum and White Crappie (Figure 5-3) than pre-flood levels. Relative
abundance of adult omnivores differed among flood periods (F = 13.95, df = 1, 69, p <
0.01), driven by increased relative abundance of Common Carp, Freshwater Drum,
Bluegill, and Channel Catfish (Figure 5-4).
Relative abundance of juvenile piscivores fluctuated moderately over time (F =
2.09, df = 2, 69, p = 0.05), caused by a decline in Smallmouth Bass CPUE during the
flood followed by a return to pre-flood levels after the disturbance (Figure 5-3). Adult
piscivores exhibited inconsistent CPUE among flood periods (F = 10.58, df = 1, 169, p <
132 0.01) as relative abundance of Largemouth Bass and Sauger increased after the flood
(Figure 5-4).
Aquatic Habitats
Mean main channel frequency per transect declined from 3.50 ± 0.26 before the
flood to 2.63 ± 0.19 after the disturbance (F1,59 = 7.00, P = 0.01; Figure 5-5). Similarly,
mean backwater frequency per transect decreased from 1.93 ± 0.28 before the disturbance
to 1.03 ± 0.14 afterwards (F1,59 = 8.30, P = 0.01; Figure 5-5). In contrast, mean sandbar
frequency per transect increased from 0.03 ± 0.03 before the flood to 1.40 ± 0.18 after the
flood (F1,59 = 28.02, P < 0.01; Figure 5-5). Although main channel width was stable
before and after the flood when expressed as a mean (F1,59 = 0.61, P = 0.44), it decreased
along a majority of transects (76.67%, n = 23). Mean largest-backwater area decreased
from 225,200 m2 to 186,700 m2 after the flood but the trend was not statistically
significant (F1,47 = 0.35, P = 0.56).
Discussion
The Lewis and Clark Delta fish community exhibited large-scale structural
stability and absence of an immediate flood response. Whereas species richness and
diversity often fluctuate after flooding in streams (Harrel, 1978; Cochran and Stagg,
2011; Kano et al., 2011), structural indices are more stable in channelized systems
(Jurajda et al., 2006) like the Missouri River. Our results fill an important knowledge gap
as freshwater deltas have received comparatively little flood research. Overall, the Delta
fish community exhibited short-term structural resilience to disturbance. Flood resistance
is not uncommon (Lojkasek et al., 2005; Franssen et al., 2006; Jurajda et al., 2006; Pires
133 et al., 2008) and may reflect evolutionary adaptions for persistence during and after
disturbance (Bischoff and Wolter, 2001; DeBoer et al., 2011).
Multivariate ordination revealed large-scale changes in relative abundance of
Delta fishes resulting from the 2011 flood that were subsequently examined using a
species-specific approach. A majority of Delta fish species exhibited higher relative
abundance after the 2011 flood than before. Adult introduced, native, and sport species
were more abundant after the disturbance. Common Carp, the only introduced species
sampled effectively in this study, is naturalized and highly adaptable (Penne and Pierce,
2008), so post-flood elevation in relative abundance is not surprising. Most juvenile
individuals also exhibited higher mean CPUE after the disturbance. Only juvenile
Freshwater Drum declined from pre-flood to flood and White Crappie from pre-flood to
post-flood. Small-bodied native species (i.e., Emerald Shiner, Red Shiner, and Spotfin
Shiner), displayed stable relative abundance before and after the disturbance, suggesting
flood resilience. Previous research suggests the Delta is a shallow rearing environment
with diverse aquatic habitats that support high densities of juvenile fishes under normal
hydrological conditions (Kaemingk et al., 2007; Schreck, 2010). Although larger-bodied,
adult fishes frequent the Delta during certain times of year (e.g., spawning), a universal
increase in relative abundance suggests movement from mainstem environments, which
has been observed elsewhere (Gerking, 1950; Trepanier et al., 1996). Small-bodied and
juvenile individuals likely occupied the Delta in high density and mainstem habitats in
low density before the flood, thus stable relative abundance is not surprising. Overall, our
results indicate the Delta served as both a rearing environment and local refuge for
juvenile and adult fishes during and after catastrophic disturbance, supporting previous
134 (Kaemingk et al., 2007; Graeb et al., 2009) and concurrent (Carlson et al. 2015) research.
Although juvenile fishes generally exhibited higher relative abundance post-flood
than pre-flood, individual species responded distinctly. Whereas juvenile River
Carpsucker exhibited the greatest post-flood increase in relative abundance, juvenile
Freshwater Drum and White Crappie declined significantly. In a study of larval fish
abundance in relation to spring discharge in the Milk River, a tributary of the Upper
Missouri River in Montana, River Carpsucker abundance peaked more than one month
before Freshwater Drum in two consecutive years (Bednarski et al., 2008). Although we
captured age-0 rather than larval River Carpsucker and Freshwater Drum in MF
sampling, species-specific phenology may partially explain differences in relative
abundance as River Carpsucker spawn earlier than Freshwater Drum. However,
fluctuation in CPUE among flood periods may reflect discharge and its associated effects
on spawning success (Koel and Sparks, 2002) more than season as later peaks in
discharge benefit some species and earlier peaks others (Bednarski et al., 2008).
Sampling during the flood was crucial for identifying potential effects on
spawning and recruitment of Delta fishes. Although relative abundance of juvenile
piscivores differed among flood periods in multivariate space, significance was
attributable solely to Smallmouth Bass. Juvenile Smallmouth Bass were less abundant
during the flood than before or after the disturbance, suggesting high discharges impaired
spawning. Effects of high discharges on Smallmouth Bass spawning and recruitment are
well-understood and nearly universally negative (Smith et al., 2005). Floods scour nests
(Montgomery and Fickeisen, 1980; Lukas and Orth, 1995), displace fry (Harvey, 1987;
Simonson and Swensen, 1990; Sabo and Orth, 1994), and decrease availability of shallow
135 pools (Aadland, 1993). These effects reduce recruitment when high flows occur during or
immediately after spawning (Smith et al., 2005), as in the Delta in 2011.
A trophic approach to flood assessment in the Delta was informative. Although
the Delta’s role as a refuge environment explained observed fluctuations in relative
abundance, trophic partitioning enhanced resolution of flood effects. Post-flood relative
abundances increased for five omnivores (Bluegill, Black Crappie, Channel Catfish,
Common Carp, Freshwater Drum) and two piscivores (Largemouth Bass, Sauger) but
only one invertivore (Shorthead Redhorse), suggesting differential flood response among
trophic groups. Although relative abundance of small-bodied invertivores likely remained
stable after the flood due to pre-flood occupation of the Delta and use of slackwater
refugia (e.g., submerged structures, low velocity channel margins, backwater pools;
Franssen et al., 2006), a trophic explanation (e.g., stable or reduced invertebrate density)
is plausible. Similarly, omnivore abundance may have increased due to large diet breadth
and use of a post-flood resource pulse. However, comprehensive diet analyses would be
necessary to verify trophic explanations.
Our assessment of changes in aquatic habitats in the Delta after the flood provided
insight for interpreting fish community trends. Channels narrowed and became less
numerous (along with backwaters), whereas sandbars became more numerous in the
Delta after the flood, suggesting an overall reduction in habitat complexity and riparian
vegetation removal. A decline in side channel and backwater abundance had potential to
diminish spawning and rearing habitats in the Delta, but a concomitant increase in
sandbars and associated shallow-water nursery habitat may have compensated for
potential losses. The Delta functioned as a shallow-water refuge environment for a
136 resilient fish community during the flood. Resilience in relative abundance may permit
resilience in reproduction and recruitment given that availability of shallow-water habitat
is still higher in the Delta compared to main channels depauperate in slackwater habitat
necessary for spawning. Future research exploring the association between recruitment
and aquatic habitat in the Delta would provide a more comprehensive picture of the
effects of post-flood habitat changes.
We conclude the Lewis and Clark Delta acts as a refuge environment for fishes
during flood and non-flood periods. The Delta fish community expanded numerically and
stabilized structurally as a majority of species across trophic guilds and age classes
became more abundant after the flood. Under a stable hydrological regime, the Delta
contains habitats that simulate those of the historic Missouri River and support high
species richness and diversity (Kaemingk et al., 2007; Graeb et al., 2009). Our results
indicate the Delta’s role as a refuge is amplified during and immediately after flooding.
We recommend continued assessment and monitoring to describe long-term disturbance
effects in the Delta.
Acknowledgments
We thank L. Heironimus, C. Hayer, T. Rapp, and J. Mecham from South Dakota
State University for field assistance. We thank B. Hanten, H. Meyer, K. Potter, G.
Knecht, J. Sorensen, and R. Trible from the South Dakota Department of Game, Fish and
Parks for technical advice and assistance. Funding for this project was provided by the
Federal Aid in Sport Fish Restoration program, Project 3M3328, Study 1526,
administered by the South Dakota Department of Game, Fish and Parks, United States
137 Fish & Wildlife Service, South Dakota State University Department of Natural Resource
Management, and South Dakota Agricultural Experiment Station.
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145 Tables
Table 5-1. Fish species collected from 2004–2012 in the Lewis and Clark Delta organized
by introduction history (SBN = small-bodied native) and trophic guild. Sample sizes of
fishes captured by electrofishing (EF) are listed by length category (adult, juvenile/smallbodied). Only juvenile fishes were sampled with mini-fyke netting (MF).
146 Table 5-2. Sampling distribution of fishes collected during flood periods (pre-flood
[2004-2010], flood [2011], post-flood [2012]) in the Lewis and Clark Delta. Fishes are
organized by introduction history, trophic guild, and gear type. Electrofishing (EF) data
included adults in all categories and guilds except small-bodied native (SBN),
invertivore, and planktivore, which represent juvenile and small-bodied individuals. EF
was not conducted during the flood. Mini-fyke net (MF) data included juvenile
individuals in all categories and guilds except those mentioned above.
Category
Time
EF
MF
172
60
Flood
n/a
31
Post
105
68
277
233
n/a
189
422
504
n/a
261
765
1651
n/a
257
1908
91
n/a
46
137
1637
n/a
278
1915
363
n/a
283
646
357
n/a
159
516
159
3641
382
489
4512
4824
555
446
5821
n/a
n/a
n/a
n/a
85
16
309
410
n/a
n/a
n/a
n/a
1019
405
319
1743
922
115
113
1150
Introduced Pre
Total
Native
Pre
Flood
Post
Total
Sport
Pre
Flood
Post
Total
SBN
Pre
Flood
Post
Total
Detritivore Pre
Flood
Post
Total
Invertivore Pre
Flood
Post
Total
Omnivore Pre
Flood
Post
Total
Piscivore Pre
Flood
Post
Total
147 Table 5-3. Structural indices (species richness, evenness [J’], diversity [Fisher’s α]) of
fishes collected during flood periods (pre-flood [2005, 2008-2009], post-flood [2012]) in
the Lewis and Clark Delta. The entire species pool was reduced to common species (>
1% of total catch during a particular sampling event) to remove effects of rare species
and species difficult to collect. Letters indicate significant differences among flood
periods. Year
Flood
Richness
J’
Fisher's α
z
0.882
x
0.796
y
0.725
y
0.835
2005
Pre
25
2008
Pre
15
2009
Pre
22
2012
Post
21
z
4.56
z
y
2.17
x
3.48
y
3.27
w
y
x
148 Figures
Figure 5-1. Missouri River watershed and location of the Lewis and Clark Delta along the
South Dakota-Nebraska border.
149 Figure 5-2. Gauge height in the Lewis and Clark Delta, South Dakota from 1980-2012.
Flooding in 2011 represented the most severe hydrological disturbance in recent history.
Gauge height was measured at a United States Geological Survey site in Springfield, SD
(USGS 06466700, Latitude 42°51'21", Longitude 97°53'06").
150 Figure 5-3. Nonnetric multidimensional scaling (NMS) ordinations of catch-per-uniteffort (CPUE) for juvenile native and sport fish sampled during pre-flood (2004-2010),
flood (2011), and post-flood (2012) periods (symbols) in the Lewis and Clark Delta,
South Dakota. Multivariate heterogeneity is explained by directional changes in CPUE of
fishes categorized by introduction history (Introduced, Native, Sport), trophic guild (D =
detritivore; O = omnivore; P = piscivore), and species (CAP = common carp; FRD =
freshwater drum; RIC = river carpsucker; BGL = bluegill; BLC = black crappie; SMB =
smallmouth bass; WHC = white crappie). NMS was performed in Primer (PRIMER-E
Ltd, Plymouth, United Kingdom; Clarke and Gorley 2006) using Bray-Curtis ecological
distance. Sample sizes are included in Table 5-1.
151 Figure 5-4. Nonmetric multidimensional scaling (NMS) ordinations of catch-per-uniteffort (CPUE) for adult native and sport fish and small-bodied native species sampled
during pre-flood (2004-2010), flood (2011), and post-flood (2012) periods (symbols) in
the Lewis and Clark Delta, South Dakota. Multivariate heterogeneity is explained by
directional changes in CPUE of fishes categorized by introduction history (Introduced,
Native, Sport, Small-bodied Native [SBN]), trophic guild (D = detritivore; I =
Introduced; O = omnivore; P = piscivore), and species (CAP = common carp; FRD =
freshwater drum; RIC = river carpsucker; SHR = shorthead redhorse; BGL = bluegill;
CCF = channel catfish; LMB = largemouth bass; SAR = sauger; EMS = emerald shiner;
RES = red shiner; SFS = spotfin shiner). NMS was performed in Primer (PRIMER-E Ltd,
Plymouth, United Kingdom; Clarke and Gorley 2006) using Bray-Curtis ecological
distance. Sample sizes are included in Table 5-1.
152 Figure 5-5. Pre-flood (June 2010) and post-flood (October 2011) frequency of channels,
backwaters, and sandbars in the Lewis & Clark Delta by equidistant transect between the
Niobrara River confluence and downstream limit of the Delta.
153 CHAPTER 6: SUMMARY, MANAGEMENT RECOMMENDATIONS, AND
RESEARCH NEEDS
Summary
A catastrophic flood in the Missouri River in 2011 exerted effects on fish
populations and aquatic habitats at statewide and local scales in South Dakota. We
assessed natal origins, movement, and entrainment of walleye Sander vitreus in four
Missouri River reservoirs before, during, and after this disturbance using otolith
microchemistry. We also evaluated how the flood affected the Lewis & Clark Delta fish
community and aquatic habitats. These analyses furnished a spatially and temporally
robust investigation of a major disturbance.
Overall, otolith microchemistry was an effective, high-resolution tool for studying
natal origins, movement, and entrainment of walleye in Missouri River reservoirs. Spatial
variability and temporal stability in water chemistry and positive, proportional waterotolith relationships laid a foundation for informative microchemistry analyses. K-nearest
neighbor discriminant analysis indicated that bivariate chemical signatures yielded the
highest natal reclassification accuracies in tributaries, reflecting heterogeneous surficial
and bedrock geology in these sites. Lake Oahe tributaries, particularly the Moreau and
Cannonball rivers, were natal sites for a large portion of adults and the majority of
tributary-natal adults. In addition, walleye natal contribution was high in the Lewis &
Clark Delta, where previous researchers (Kaemingk et al. 2007; Graeb et al. 2009) have
linked quality of spawning and nursery habitat to high species richness and diversity.
154 Walleye movement within and among Missouri River reservoirs was dominated
by downstream passage and site residency, whereas upstream movement was relatively
uncommon. Downstream movement was pervasive during the flood, yet movement
patterns were not universally similar among reservoirs before and after the disturbance.
Downstream movement from North Dakota to South Dakota was more extensive than
upstream movement from South Dakota to North Dakota. The percentage of entrained
walleye progressively increased moving downstream from Lake Sharpe (18.87%) to Lake
Francis Case (24.39%) to Lewis & Clark Lake (34.15%), as did percent floodentrainment. Nearly two-thirds (64.71%) of entrained walleye moved through dams
during the flood year. Otolith microchemistry successfully predicted entrainment during
floods and normal hydrological regimes. It also illuminated connectivity of walleye
populations and may facilitate holistic riverscape management in Missouri River
reservoirs.
Catastrophic flooding in the Missouri River in 2011 also provided an opportunity
to measure immediate ecological effects of a large disturbance in the Lewis & Clark
Delta. Community evenness (J’) consistently decreased from 0.882 to 0.725 before the
flood but rose to 0.835 after the disturbance. Pre-flood species richness declined from 25
to 15, whereas post-flood richness stabilized above 20. Diversity (Fisher’s α) decreased
from 4.56 to 3.48 before the flood and stabilized at 3.27 after the disturbance, indicating
absence of an immediate structural flood response. Nonmetric multidimensional scaling
permitted multivariate assessment of flood effects on relative abundance of specific
segments of the Delta fish community. A majority of species were more abundant after
the flood regardless of native/nonnative status, trophic guild, and length. Relative
155 abundance of juvenile freshwater drum Aplodinotus grunniens and white crappie
Pomoxis annularis declined from pre-flood levels, whereas relative abundance of smallbodied native species remained stable. Overall, the Delta served as a refuge environment
for juvenile and adult fishes during and immediately after catastrophic disturbance.
Management Recommendations
Overview
Management implications of our research are abundant and diverse. Riverscape
walleye management programs are imperative as important natal tributaries and
embayments are common throughout reservoirs and individuals move extensively within
and among impoundments. We recommend that state agencies collaborate to use our
results to design and implement large-scale management strategies, particularly in
interjurisdictional fisheries such as Lake Oahe. For example, we advise managers to
focus egg take operations in sites where high numbers of walleye reproduce to streamline
spawning operations, conserve areas with fewer spawners, and promote spatially
diversified reproduction. For habitat conservation, we suggest managers protect natal
sites from sedimentation, shoreline erosion, damming, pollution, and other stressors to
promote riparian functions and services (e.g., sediment control, nutrient cycling, flood
control) at impoundment and riverscape scales. For habitat restoration, we recommend
managers enhance physical and chemical habitats in degraded natal areas by dredging
sediments, stabilizing shorelines, planting riparian buffers, and/or mitigating point-source
pollution. We advise managers to educate and incentivize landowners throughout the
riverscape to minimize nutrient and sediment delivery into reservoirs. Moreover, we
156 suggest managers ensure water level manipulations maintain or improve (rather than
degrade) walleye spawning and nursery habitats.
Natal Origins
Management application: protect tributaries and non-tributary sites
Recommendations: Our results indicated that tributaries are important for walleye
reproduction, but not all individuals spawn in them as embayments and mainstem sites
also function as natal environments. In all reservoirs but Lake Oahe, the majority of adult
walleyes spawned in non-tributary habitats. Although assignment accuracy to these natal
sites was low due to relative spatial homogeneity in trace element signatures, it is critical
for managers to recognize the importance of embayments and mainstem sites as
reproductive habitats. We recommend managers integrate anecdotal information on
spawning locations with future research (e.g., telemetry, otolith microchemistry with
novel signatures) to quantify the importance of non-tributary spawning sites. This will
foster integrative management programs in which tributaries and other critical natal
environments are prioritized for egg take, habitat protection and restoration, harvest
regulations, and other management actions.
Management application: distinguish spawning and recruitment
Recommendations: Although we used otolith microchemistry to identify important
locations for hatching and recruitment, our results may not be representative of spawning
if survival of age-0 walleyes varies among sites. Thus, we recommend managers use
predictive modeling based on existing and future population data to link spawning with
recruitment. This will enable managers to identify spawning sites that function as
157 “recruitment hotspots” and design corresponding management strategies (e.g., minimum
size limits) that protect recruits in these areas.
Movement
Management application: recognize intra-reservoir and interjurisidictional movement
Recommendations: Because North Dakota and South Dakota natal sites contribute
walleyes to the opposite state and individuals move extensively between states,
collaborative interstate management is necessary. We recommend that managers ensure
that harvest regulations (i.e., creel limits, slot limits, one-over rules) are compatible
among states under normal hydrological conditions and after floods. In addition, we
advise managers to collaborate on habitat protection and restoration activities (e.g.,
riparian zone preservation/enhancement, sediment dredging), particularly at important
spawning locations (e.g., Moreau and Cannonball rivers). Moreover, we recommend
managers in both states jointly conduct human dimensions research to understand
stakeholders in both states and use this information to design riverscape-scale
management strategies and public engagement initiatives.
Entrainment
Management application: distinguish baseline and flood-induced entrainment and
understand spatial patterns
Recommendations: Although walleye entrainment through mainstem Missouri River
dams increased during the flood, fish passage also occurred during non-flood years. Such
baseline entrainment has important management implications. First, walleye populations
in Missouri River reservoirs are more connected than previously believed. That upper
158 reservoirs subsidize populations in lower impoundments indicates the importance of
riverscape management. For example, large-scale habitat protection and restoration
programs are likely to enhance walleye populations inside and outside the specific
reservoirs where they are implemented. In addition, entrainment increased during the
flood and moving downstream in Missouri River reservoirs, potentially necessitating
harvest regulations that account for population subsidization in lower reservoirs. Much
like baseline entrainment, the fact that lower reservoirs, with fewer important natal sites,
contain higher proportions of entrained walleye signifies the significance of habitat
conservation in upper reservoirs, where natal environments are more abundant.
Management application: conduct cost-benefit analysis of entrainment mitigation
Recommendations: Given that managers can mitigate entrainment using established
methods (e.g., strobe lights, sound deterrents), we project potential outcomes and
implications under different management scenarios. Currently in the absence of
entrainment mitigation, the Missouri River functions as a riverscape of connected
impoundments with interconnected walleye populations. Entrainment is known to occur
under all hydrological conditions, and it increases with floods and moving downstream.
Under these conditions, we recommend managers continue to monitor baseline
entrainment, which will inform future management strategies in diverse hydrological
regimes. In addition, we advise managers to quantify site-specific spawning-recruitment
relationships and protect populations via habitat conservation, particularly in upstream
reservoirs that subsidize lower ones. In contrast, if managers attempted to mitigate
entrainment, they would rely solely on natural reproduction to replenish populations in
lower reservoirs. Upper reservoirs with abundant natal habitats (i.e., Oahe) may be
159 relatively self-sustaining, but lower reservoirs with fewer natal environments and high
proportions of entrained individuals may not be. Our results suggest that entrainment
plays a vital role in structuring walleye populations in Missouri River reservoirs. For
example, 47.5% of adult walleye in Lake Francis Case hatched in upstream reservoirs, as
did 35% of adults in Lake Lewis & Clark. Although these reservoirs contain natal sites,
our results suggest natural reproduction may not match entrainment subsidization. We
recommend managers compare the advantages and disadvantages of extant entrainment
in relation to goals, which will promote science-based walleye management in Missouri
River reservoirs.
Research Needs
This project contributed to scientific understanding of flood effects on fishes and
aquatic habitats in the Missouri River. Additional research would expand the depth and
breadth of this growing body of knowledge. In particular, otolith microchemistry studies
exploring natal origins, movement, and entrainment of other species (e.g., gizzard shad
Dorosoma cepedianum, rainbow smelt Osmerus mordax, lake herring Coregonus artedi,
smallmouth bass Micropterus dolomieu, northern pike Esox lucius) would broaden our
understanding of fish environmental history in Missouri River reservoirs and yield
additional information germane for management. Microchemistry conducted at finer
temporal scales (e.g., interannually, monthly, daily) would illuminate fish movement
patterns that were not discernible in the present study. The relative importance of
Missouri River embayments to recruitment of predator and prey populations was only
marginally detectable using bivariate (e.g., Sr:Ca, Ba:Ca) signatures. Additional chemical
tracers (e.g., oxygen and carbon isotopes) may provide higher resolution for assessing
160 natal origins and movement in these locations. Future researchers could also explore
advanced statistical techniques (e.g., Bayesian hierarchical mixture models; Pflugeisen
and Calder 2013) to analyze otolith microchemistry data and advance fisheries
management in Missouri River reservoirs. Finally, long-term post-flood responses of the
Lewis & Clark Delta fish community and aquatic habitats would complement existing
knowledge and facilitate science-based sediment and water management in the region.
161 APPENDICES
162 Appendix A. Reservoir, site, date collected, total length (mm), date ablated, terminal
otolith Sr:Ca, and terminal otolith Ba:Ca of age-0 walleye collected in fall 2012 in
tributaries and at embayment and mainstem sites of South Dakota Missouri River
reservoirs. “Am. Crk. Emb.” and “GPD” are abbreviations for American Creek
Embayment and Gavin’s Point Dam, respectively. Only Sr:Ca and Ba:Ca are reported
here as only these elements had sufficient resolution to be used to evaluate walleye
environmental history. Age-0 otoliths with outlier chemical concentrations were removed
from this table and all analyses reported above. SE denotes standard error.
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Site
Grand R.
Grand R.
Grand R.
Grand R.
Grand R.
Moreau R.
Cheyenne R.
Cheyenne R.
Cheyenne R.
Cheyenne R.
Knife R.
Heart R.
Cannonball R.
Cannonball R.
Beaver Bay
Beaver Bay
Beaver Bay
Beaver Bay
Beaver Bay
W. Pollock
W. Pollock
W. Pollock
Swan Creek
Swan Creek
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
Minneconjou
Minneconjou
Minneconjou
Minneconjou
Okobojo
Okobojo
Okobojo
Okobojo
Okobojo
Okobojo
Okobojo
Bad R.
Bad R.
Bad R.
Bad R.
Stilling Basin
Stilling Basin
Stilling Basin
Stilling Basin
Stilling Basin
Ft. George
Ft. George
Ft. George
Ft. George
Date Collected Total Length (mm) Sr:Ca (μmol/mol)
10/23/2012
151
1.385839108
10/23/2012
162
1.325829887
10/23/2012
135
1.296864413
10/23/2012
150
1.328675214
10/23/2012
132
1.535023311
10/30/2012
161
1.182198695
11/7/2012
191
1.486891533
11/7/2012
157
1.429199537
11/7/2012
158
1.474902186
11/7/2012
146
1.524458516
10/23/2012
194
1.721561488
10/23/2012
139
1.517304182
10/23/2012
138
1.941737641
10/23/2012
127
2.043973623
9/6/2012
100
1.265681427
9/6/2012
90
1.108033244
9/6/2012
130
1.314305779
9/6/2012
100
1.246236724
9/6/2012
110
1.325312588
10/24/2012
124
1.442610596
10/24/2012
109
1.405988168
10/24/2012
136
1.273363695
10/30/2012
114
1.73336167
10/30/2012
139
1.69787257
10/10/2012
177
1.131811266
10/10/2012
147
1.220520537
10/10/2012
148
1.14384996
10/10/2012
212
1.342542075
10/10/2012
137
1.359441881
10/12/2012
152
1.349002659
10/12/2012
132
1.368535642
10/12/2012
167
1.452989094
10/12/2012
141
1.306964887
10/10/2012
153
1.385095078
10/10/2012
137
1.210672153
10/10/2012
176
1.43087336
10/10/2012
162
1.428568599
10/10/2012
170
1.380397621
10/10/2012
168
1.440847475
10/10/2012
165
1.429475346
10/16/2012
137
1.248554065
10/16/2012
127
1.214626488
10/16/2012
103
1.298525949
10/16/2012
?
1.337591671
10/9/2012
98
1.530137067
10/9/2012
135
1.386631726
10/9/2012
97
1.487620665
10/9/2012
127
1.484333819
10/9/2012
105
1.516800434
10/10/2012
138
0.903549226
10/10/2012
164
1.100905454
10/10/2012
100
1.013882909
10/10/2012
118
1.207259535
Sr:Ca SE
0.047768013
0.045690045
0.044707737
0.052742498
0.06094101
0.038284942
0.048171024
0.046281961
0.047780615
0.049380018
0.071129329
0.051319416
0.084969834
0.089453205
0.059782388
0.052339499
0.062074445
0.058863046
0.062590782
0.055059743
0.05364925
0.048599168
0.061671441
0.060412069
0.036660353
0.04128222
0.038725694
0.043498746
0.045992272
0.043675058
0.044317342
0.047050178
0.042340126
0.057238456
0.050022259
0.059127519
0.059026764
0.057036957
0.059543107
0.059102327
0.045249239
0.04401505
0.047062734
0.048473231
0.055450152
0.05024895
0.053901125
0.057528121
0.058774899
0.031018339
0.037793762
0.034796458
0.041433351
Ba:Ca (μmol/mol)
0.003696857
0.003680678
0.003397549
0.004327827
0.006398713
0.00284747
0.003187225
0.003057795
0.002734218
0.003834377
0.005711114
0.004481526
0.010265445
0.011964219
0.005468432
0.006989239
0.008696101
0.014674165
0.010540483
0.023200391
0.010847882
0.010540485
0.009804348
N/A
0.005581689
0.004481526
0.00295263
0.008922612
0.003858643
0.004311652
0.003454176
0.003640232
0.003567427
0.003405636
0.004643314
0.003365189
0.003583603
0.003429904
0.002887915
0.003446083
0.004546241
0.007223832
0.004319738
N/A
0.008963052
0.008437242
0.008380616
0.008793175
0.008235007
0.003599782
0.00376157
0.004926444
0.004287382
Ba:Ca SE
0.000145609
0.000145609
0.00013752
0.000177967
0.000258861
0.000121341
0.00013752
0.00012943
0.000113252
0.000161788
0.000436827
0.000283129
0.000736135
0.000857476
0.000242682
0.000315486
0.000388291
0.000647152
0.000469185
0.001003085
0.000469185
0.000453006
0.000380202
N/A
0.000234593
0.000283129
0.000186056
0.000380202
0.000242682
0.000177967
0.000145609
0.000153699
0.000153699
0.000258861
0.000355933
0.000258861
0.000275039
0.00026695
0.000218414
0.00026695
0.000153699
0.000242682
0.000145609
N/A
0.000299308
0.000283129
0.000283129
0.000380202
0.000355934
0.000258861
0.00027504
0.000355934
0.000307397
163 Appendix A, continued.
Reservoir
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
Site
Date Collected Total Length (mm) Sr:Ca (μmol/mol)
W. Bend
10/11/2012
140
0.972940563
W. Bend
10/11/2012
153
0.968003824
W. Bend
10/11/2012
157
0.900879287
W. Bend
10/11/2012
139
0.874004285
W. Bend
10/11/2012
140
0.8563227
W. Bend
10/11/2012
172
0.830681882
N. Shore
10/11/2012
126
0.843867574
N. Shore
10/11/2012
157
0.923485089
N. Shore
10/11/2012
184
0.859370447
N. Shore
10/11/2012
191
0.871397519
N. Shore
10/11/2012
153
0.890527386
N. Shore
10/11/2012
160
0.914002086
N. Shore
10/11/2012
170
0.852960308
White R.
10/2/2012
140
1.135009342
White R.
10/2/2012
163
1.143673559
FC St. B
10/1/2012
147
1.15704818
FC St. B
10/1/2012
184
1.007825172
FC St. B
10/1/2012
152
1.039964349
FC St. B
10/1/2012
158
1.199829052
FC St. B
10/1/2012
180
1.05976168
FC St. B
10/1/2012
151
1.070869342
Am. Crk. Emb.
10/1/2012
160
1.278595559
Am. Crk. Emb.
10/1/2012
165
1.227732244
Platte Crk.
10/4/2012
130
1.005344208
Platte Crk.
10/4/2012
183
1.269724306
Platte Crk.
10/4/2012
183
1.19611409
Platte Crk.
10/4/2012
114
1.17795394
N. Bay
10/4/2012
143
1.363182216
N. Bay
10/4/2012
139
1.290037881
N. Bay
10/4/2012
138
1.14916451
N. Bay
10/4/2012
127
1.27984956
N. Bay
10/4/2012
140
1.228505147
N. Bay
10/4/2012
160
1.174389916
N. Bay
10/4/2012
130
1.171808112
N. Bay
10/4/2012
151
1.237018504
Niobrara R.
10/22/2012
152
1.506712863
Niobrara R.
10/22/2012
160
1.546811272
Niobrara R.
10/22/2012
185
1.588370554
Tailrace
10/4/2012
176
1.396669138
Tailrace
10/4/2012
145
1.42241049
Tailrace
10/4/2012
149
1.509042818
Tailrace
10/4/2012
132
1.597463344
Springfield
10/11/2012
133
1.136318738
Springfield
10/11/2012
163
1.723728141
Springfield
10/11/2012
184
1.699283198
GPD
10/11/2012
156
1.730755037
GPD
10/11/2012
151
1.662471755
GPD
10/11/2012
121
1.519042116
GPD
10/11/2012
165
1.684208776
GPD
10/11/2012
154
1.633808702
Sr:Ca SE
0.042579373
0.04236528
0.039430943
0.033864518
0.033171863
0.032189553
0.028940375
0.031673212
0.029481905
0.0298975
0.03053978
0.031358372
0.029267814
0.049317022
0.05483306
0.048057642
0.041861531
0.043196465
0.049833357
0.04744055
0.04798208
0.052667534
0.050579909
0.041773375
0.055147912
0.051923923
0.051130518
0.061029175
0.057767401
0.051445353
0.057301434
0.038851637
0.037138891
0.037050732
0.039116106
0.064920635
0.066645975
0.068472065
0.053296638
0.061293643
0.057603687
0.060966211
0.045967077
0.077615119
0.068736522
0.06112992
0.058737109
0.058674149
0.065008791
0.063056764
Ba:Ca (μmol/mol)
0.004117504
0.003308564
0.003543156
0.004360185
0.00362405
0.004004252
0.002984988
0.003721123
0.002904094
0.003704945
0.0028232
0.00364023
0.005063964
N/A
0.004384453
0.00537945
0.002766574
0.003284296
0.003340921
0.004060878
0.004271202
0.003575514
0.002701859
0.001981902
0.004530063
0.00497498
0.002855558
0.002984988
0.003518888
0.002604786
0.003381368
0.004206488
0.005500791
0.00443299
0.003243849
0.007579766
0.031945032
0.007215743
0.003923359
0.00429547
0.003187223
0.009836709
0.006277372
0.006722291
0.013622544
0.006236925
0.006131763
0.005007337
0.008590941
0.005808188
Ba:Ca SE
0.000299308
0.000242682
0.000258861
0.000186056
0.000153699
0.000169877
0.000169877
0.000210324
0.000210324
0.00026695
0.000161788
0.000210324
0.000291218
N/A
0.000210324
0.000242682
0.00012943
0.000153699
0.000153699
0.000202235
0.000218414
0.00013752
0.000105162
0.0000971
0.000202235
0.000218414
0.00012943
0.000145609
0.000177967
0.00012943
0.000169877
0.000177967
0.000234593
0.000194146
0.00013752
0.000372112
0.001536986
0.000355934
0.000153699
0.000210324
0.00012943
0.000380202
0.000323576
0.000355934
0.000695688
0.000250771
0.000250771
0.000234593
0.00039638
0.00026695
164 Appendix B. Mean terminal otolith Sr:Ca and Ba:Ca (μmol/mol) signatures of age-0
walleye collected in tributaries (n = 9) and embayments (n = 17) in South Dakota
Missouri River reservoirs in fall 2012. These signatures were used to create a “blueprint”
of site-specific water chemistry for retrospective site assignment of adult walleye, which
laid a foundation for evaluating entrainment through mainstem dams before, during, and
after the 2011 flood. “Am. Crk. Emb.” and “GPD” are abbreviations for American Creek
Embayment and Gavin’s Point Dam, respectively. SE denotes standard error.
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Site
Sr:Ca (μmol/mol) Sr:Ca SE Ba:Ca (μmol/mol) Ba:Ca SE
Grand R.
1.3744
0.0427
0.0043
0.0005
Moreau R.
1.1822
0.0028
Cheyenne R.
1.4789
0.0196
0.0034
0.0002
Knife R.
1.7216
0.0057
Heart R.
1.5173
0.0045
Cannonball R.
1.9929
0.0511
0.0115
0.0006
Beaver Bay
1.2519
0.0389
0.0109
0.0007
W. Pollock
1.3740
0.0514
0.0108
0.0002
Swan Creek
1.7156
0.0177
0.0098
0.0004
W. Whitlocks
1.2396
0.0480
0.0054
0.0008
Minneconjou
1.3694
0.0307
0.0039
0.0005
Okobojo
1.3866
0.0307
0.0035
0.0005
Bad R.
1.2748
0.0271
0.0037
0.0006
Stilling Basin
1.4811
0.0252
0.0086
0.0004
Ft. George
1.0564
0.0645
0.0041
0.0006
W. Bend
0.9005
0.0240
0.0038
0.0005
N. Shore
0.8794
0.0116
0.0032
0.0002
White R.
1.0345
0.0278
0.0039
0.0005
Stilling Basin
1.0892
0.0301
0.0035
0.0006
Am. Crk. Emb.
1.2532
0.0254
0.0033
0.0004
Platte Crk.
1.1623
0.0559
0.0046
0.0005
N. Bay
1.2367
0.0255
0.0033
0.0004
Niobrara R.
1.5473
0.0236
0.0074
0.0003
Tailrace
1.4814
0.0455
0.0034
0.0002
Springfield
1.5198
0.1919
0.0065
0.0002
GPD
1.6461
0.0355
0.0059
0.0005
165 Appendix C. Ablation period, reservoir, site, total length (mm), otolith radius (μm), and
age of adult walleye 2013 in embayment and mainstem sites of South Dakota Missouri
River reservoirs. GPD and ODF are abbreviations for Gavin’s Point Dam and Oahe
Dame Face, respectively. Only Sr:Ca and Ba:Ca are reported here as only these elements
had sufficient resolution to be used to evaluate walleye environmental history. Age-0
otoliths with outlier chemical concentrations were removed from this table and all
analyses reported above. SE denotes standard error.
Ablation Period
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
October 2013
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Site
Bush
Bush
Bush
Bush
Bush
Bush
Bush
Bush
Bush
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
ODF
Sutton
Sutton
Sutton
Joe Crk.
Joe Crk.
Joe Crk.
West Bend
West Bend
Beaver
Beaver
Beaver
Beaver
Beaver
Beaver
Beaver
Beaver
Beaver
Beaver
Total Length Otolith radius
355
N/A
310
N/A
326
368.3
632
1683.6
399
799.7
353
560.5
318
518.5
382
869.4
576
712.8
300
513.7
268
422.2
300
491.9
305
638.4
319
496.3
346
492.1
360
772.7
270
656.5
280
457.1
299
627.5
548
793.1
327
509.6
320
550.1
309
749.7
312
463.5
291
415.6
327
574
300
476.2
352
404.4
405
502.4
262
467.2
276
1875.4
246
1249.7
389
2292.2
384
926
386
723.4
N/A
1297.1
N/A
1011.8
N/A
976.5
N/A
959.5
N/A
1129
N/A
576.5
N/A
1797.4
N/A
1074.5
N/A
1031.8
N/A
1098
Age
3
2
3
6
4
3
3
4
5
2
2
2
2
3
3
3
2
2
2
6
4
3
3
2
2
3
2
3
4
2
2
2
4
3
3
4
3
8
4
4
3
7
5
4
3
166 Appendix C, continued.
Ablation Period
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Site
Cattail Bay
Cattail Bay
Cattail Bay
Cattail Bay
Cattail Bay
Cattail Bay
Cattail Bay
Cattail Bay
Cattail Bay
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Cow Creek
Fort Yates
Fort Yates
Fort Yates
Fort Yates
Fort Yates
Fort Yates
Fort Yates
Fort Yates
Fort Yates
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
Total Length Otolith radius
N/A
1032.7
N/A
1132.7
N/A
1223.2
N/A
1449.6
N/A
1776.8
N/A
1248.4
N/A
1182.8
N/A
1735.9
N/A
978.8
321
829.4
461
1031
446
1131.9
447
1266.1
348
1257.3
452
1521
395
860.9
322
601.6
321
919.9
446
953.3
357
1280.5
351
1135.9
373
1178.5
N/A
881.7
N/A
881.5
N/A
1226.7
N/A
936.3
N/A
968.1
N/A
1589.5
N/A
1208.5
N/A
1595.8
N/A
802.7
N/A
1428.7
N/A
718.3
N/A
1279.2
N/A
532.9
N/A
1035.6
N/A
1079.1
N/A
787.1
N/A
885.2
N/A
824.1
375
1142.7
351
400.5
380
990.1
380
975.2
330
932.2
Age
4
4
4
4
8
4
4
5
4
3
3
3
4
3
4
3
3
3
3
3
3
3
4
4
4
6
3
6
4
9
4
4
4
4
3
3
3
3
4
4
3
2
3
3
1
167 Appendix C, continued.
Ablation Period
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Site
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
W. Whitlocks
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
Joe Creek
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
N. Shore
Total Length Otolith radius
322
778.1
564
771.5
332
1160.9
305
736.9
330
928.5
410
898.5
306
1265.6
637
974.1
350
794.7
331
1085.6
329
955.7
372
1044.6
389
909.7
410
604.6
385
677.2
440
915.3
351
702
407
634
425
663.9
419
734
264
438.1
258
968.3
273
642.7
296
656.4
439
1009.5
400
703.6
354
414.8
335
664.4
463
439.2
317
862.6
382
302.3
415
1000.5
430
816.8
345
571.4
392
909.5
382
843.7
375
681.3
372
1026.6
358
986
365
710.1
386
609.7
440
670
410
687.1
Age
3
2
2
2
2
3
3
3
2
2
3
3
3
3
3
3
3
4
4
7
1
2
2
2
4
3
3
2
5
2
2
3
3
2
3
3
2
3
3
3
2
4
3
168 Appendix C, continued.
Ablation Period
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
Reservoir
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
Site
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
Snake Creek
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
St. Basin
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Chamberlain Area
Total Length Otolith radius
235
371.6
141
511.6
318
661.2
271
858.2
268
840.3
330
1296.3
230
693.4
218
773.5
316
798.1
242
391.2
240
422.5
312
877.6
285
1022.3
364
690.8
382
612.8
405
842.4
333
334.5
515
693.3
410
486.3
313
532.1
399
596.1
360
439.7
337
761.1
349
519.2
320
616.2
355
575.7
410
465.7
680
1327.6
355
302.4
412
459
290
525.9
364
725.4
371
892.1
355
581.4
315
424.5
349
530.8
200
314.4
333
365.2
360
719.4
352
660.1
363
540.1
353
749.3
352
938
344
854.5
368
601.4
269
463.3
362
788.1
352
582.7
Age
2
2
3
2
2
3
2
1
3
2
2
2
2
3
3
4
3
6
3
2
3
3
2
2
3
2
3
11
2
4
2
3
4
3
2
3
2
2
3
2
3
3
3
3
4
2
2
2
169 Appendix C, continued.
Ablation Period
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
Reservoir
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
FC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
Site
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
N. Point
GPD
GPD
GPD
GPD
GPD
GPD
GPD
GPD
GPD
GPD
GPD
GPD
GPD
GPD
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
LC Tailrace
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Total Length Otolith radius
339
558.4
367
674.8
384
602.3
395
633
356
660.6
278
530.2
378
428.4
369
457.5
532
969.3
361
1214.7
351
988.1
425
637.1
375
676.9
384
505.5
367
552.5
512
742.7
383
494.9
379
470.1
447
714.1
555
1209.1
321
510.5
351
354.2
479
698.5
369
506.2
324
478.4
473
704.7
283
361.4
498
1150.2
508
792.5
384
838.8
349
551.2
357
742
435
1215.8
333
1109.8
412
798.7
372
1236.4
349
912.9
330
1225.3
515
1644
465
1587.5
514
983.5
509
677.1
212
862.6
262
462.9
349
382.2
407
625.3
Age
2
3
3
3
3
2
3
2
6
4
3
3
3
2
3
7
3
3
4
10
2
3
4
3
3
5
2
5
5
4
3
3
5
3
3
2
3
2
5
6
6
7
2
2
2
5
170 Appendix C, continued.
Ablation Period
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
April 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
Reservoir
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
LC
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Site
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
Tabor
4 Mile
4 Mile
4 Mile
4 Mile
4 Mile
4 Mile
4 Mile
4 Mile
4 Mile
4 Mile
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Garrison Tailrace
Kimball
Kimball
Kimball
Kimball
Kimball
Kimball
Kimball
Kimball
Kimball
Kimball
Total Length Otolith radius
530
1173.1
279
328
448
608.2
478
748.5
443
579.1
453
668.3
422
492.8
441
771.9
290
515.3
434
527.6
428
818
368
488.9
456
628.7
349
480
481
584
N/A
605.6
N/A
1065.1
N/A
635
N/A
636.7
N/A
704.5
N/A
1151
N/A
532.2
N/A
998.7
N/A
829.7
N/A
940.1
N/A
562.5
N/A
745.4
N/A
742.4
N/A
528.4
N/A
667.7
N/A
769.4
N/A
905.5
N/A
1040.1
N/A
706.5
N/A
916.5
N/A
626.8
N/A
768.8
N/A
810.2
N/A
590.9
N/A
704.3
N/A
512.2
N/A
556.8
N/A
638.5
N/A
608.6
N/A
725.6
Age
11
2
5
8
4
6
4
5
2
4
5
3
5
2
5
3
7
5
4
4
8
3
4
4
4
3
4
4
3
3
3
4
5
4
4
3
4
5
3
3
3
3
3
3
3
171 Appendix C, continued.
Ablation Period
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Site
Kneifel
Kneifel
Kneifel
Kneifel
Kneifel
Kneifel
Kneifel
Kneifel
Kneifel
Kneifel
Langliers
Langliers
Langliers
Langliers
Langliers
Langliers
Langliers
Langliers
Langliers
Langliers
Peoria Flats
Peoria Flats
Peoria Flats
Peoria Flats
Peoria Flats
Peoria Flats
Peoria Flats
Peoria Flats
Peoria Flats
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Porcupine Creek
Stanton
Stanton
Stanton
Stanton
Stanton
Stanton
Stanton
Stanton
Stanton
Stanton
Total Length Otolith radius
N/A
559.7
N/A
644.2
N/A
820.6
N/A
619.3
N/A
764.8
N/A
531.1
N/A
726.5
N/A
680.2
N/A
745.1
N/A
531.4
N/A
737.5
N/A
1091.8
N/A
786.5
N/A
1217.3
N/A
1349
N/A
946.5
N/A
728.3
N/A
N/A
N/A
1155.8
N/A
N/A
N/A
703.3
N/A
851.8
N/A
462
N/A
728.5
N/A
781.4
N/A
802.6
N/A
440.4
N/A
723.1
N/A
663.4
N/A
508.1
N/A
759.7
N/A
730.4
N/A
879.9
N/A
980.6
N/A
406.9
N/A
615.1
N/A
854.5
N/A
667.1
N/A
684.8
N/A
423
N/A
720.3
N/A
690.5
N/A
571.2
N/A
978.2
N/A
510
N/A
519.3
N/A
620.4
N/A
580.2
N/A
458.8
Age
4
4
3
4
4
4
4
4
4
4
3
6
4
4
5
4
4
N/A
4
N/A
3
4
3
4
3
4
3
3
4
3
4
3
4
4
4
4
4
3
4
2
3
3
3
4
5
3
4
2
3
172 Appendix C, continued.
Ablation Period
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
July 2014
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Site
Steckel
Steckel
Steckel
Steckel
Steckel
Steckel
Steckel
Steckel
Steckel
Steckel
Vanderwall
Vanderwall
Vanderwall
Vanderwall
Vanderwall
Vanderwall
Vanderwall
Vanderwall
Washburn
Washburn
Washburn
Washburn
Washburn
Washburn
Washburn
Washburn
Washburn
Washburn
Total Length Otolith radius
N/A
692.1
N/A
838.3
N/A
665.4
N/A
824
N/A
806.7
N/A
902.9
N/A
797.1
N/A
801.7
N/A
685.3
N/A
720
N/A
1152.5
N/A
891.6
N/A
785.3
N/A
N/A
N/A
1163.2
N/A
792.4
N/A
874.5
N/A
1021.5
N/A
767.5
N/A
511.1
N/A
838.7
N/A
543.5
N/A
659.2
N/A
713.6
N/A
580.9
N/A
759.5
N/A
512.2
N/A
664.9
Age
4
4
4
4
4
5
8
6
4
4
3
3
4
N/A
6
3
2
3
5
3
3
4
4
5
3
4
3
3
173 Appendix D. Trace element signatures, reproducibility, and precision of standard
reference materials used for laser ablation inductively coupled plasma mass spectrometry
(LA-ICPMS) at the University of California–Davis.
174 Table D-1. Standard reference materials for age-0 walleye otoliths ablated in October
2013. USGS synthetic glass standard GSE-1G was used as the calibration standard, and
two additional reference standards (GSD-G1 and MACS-3) were used as quality controls
for verification of instrument accuracy and precision. Each standard was ablated in three
to five locations after every 4-10 samples to adjust for possible instrument drift.
Group
Oahe 1
Oahe 2
Oahe 3
Oahe 4
Oahe 5
Oahe 6
Oahe 7
Standard Ablation #
GSD
1
2
3
4
5
6
MACS-3
1
2
3
GSD
1
2
3
4
5
MACS-3
1
2
GSD
1
2
3
4
5
6
MACS-3
1
2
3
GSD
1
2
3
4
5
6
MACS-3
1
2
3
GSD
1
2
3
4
5
6
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
5
MACS-3
1
2
23
Na
26178.60
26272.97
27100.65
26800.11
26065.34
26815.52
4606.90
4581.06
4399.56
28679.14
29221.08
28224.26
27606.25
25505.45
6271.22
5530.62
27430.17
24813.31
27091.87
26679.31
25920.26
25677.94
4799.66
4950.59
4928.13
27741.67
28604.78
27253.48
28013.89
30249.57
27682.91
4850.36
5018.02
4819.90
29574.02
29690.81
26791.71
25655.51
29743.82
768.16
5656.92
6091.75
28133.79
28870.54
29378.30
25461.01
5928.63
5287.15
28679.14
29221.08
28224.26
27606.25
25505.45
6271.22
5530.62
24
Mg
21312.29
21121.27
21670.66
21128.59
20532.97
21123.56
1653.30
1626.12
1598.10
26065.24
25043.90
23438.63
22207.77
21291.56
1916.28
1671.68
21701.84
20976.13
21413.62
21697.00
21168.25
21170.64
1672.64
1714.02
1671.26
23707.01
23386.85
22640.95
21510.99
23125.27
22105.52
1738.76
1774.87
1742.44
25256.47
23483.23
23375.71
22905.43
22696.98
246.71
1892.21
1932.02
24694.44
24262.36
23944.53
20480.88
1830.28
1731.66
26065.24
25043.90
23438.63
22207.77
21291.56
1916.28
1671.68
43
Ca
50600.01
50600.01
50600.01
50600.01
50600.01
50600.01
376933.06
376933.09
376933.09
50600.00
50600.00
50600.00
50600.00
50600.00
376933.00
376933.00
50600.01
50600.01
50600.01
50600.01
50600.01
50600.01
376933.06
376933.09
376933.06
50600.00
50600.00
50600.00
50600.00
50600.00
50600.00
376933.03
376933.03
376933.03
50600.00
50600.00
50600.00
50600.00
50600.00
50600.00
376933.00
376933.00
50600.01
50600.01
50600.00
50600.00
376933.03
376933.03
50600.00
50600.00
50600.00
50600.00
50600.00
376933.00
376933.00
55
Mn
198.30
206.21
207.32
201.43
204.59
206.48
444.46
447.97
437.21
226.82
228.26
215.67
209.60
198.00
542.79
480.95
208.87
202.48
200.59
204.64
200.48
198.12
453.20
465.58
453.15
210.53
206.43
211.30
218.49
217.06
209.43
461.69
461.52
456.20
233.62
219.67
211.20
210.18
219.20
66.75
507.52
518.82
221.35
219.32
224.78
195.51
513.05
492.64
226.82
228.26
215.67
209.60
198.00
542.79
480.95
86
Sr
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
88
Sr
68.00
70.89
70.43
69.39
70.61
70.27
7674.89
7409.53
7469.29
73.72
72.03
71.55
68.56
68.29
7881.43
7531.69
71.02
69.11
69.65
68.40
70.08
70.43
7250.08
7347.00
7303.75
70.25
69.95
71.69
68.14
66.60
66.26
7761.22
7251.27
7221.53
67.76
69.18
67.96
69.52
70.43
989.33
7667.10
7834.84
70.78
69.02
71.06
68.03
7708.34
7260.02
73.72
72.03
71.55
68.56
68.29
7881.43
7531.69
137
Ba
64.74
67.96
69.43
68.76
67.49
66.75
67.82
67.77
66.97
72.25
75.18
74.63
68.26
66.37
72.94
66.38
69.06
66.87
69.01
66.34
67.86
68.02
66.05
71.51
70.02
71.96
71.42
72.63
67.64
68.39
65.03
69.55
72.05
66.92
69.10
69.07
69.14
70.46
67.03
9.30
66.98
72.00
75.11
72.36
72.81
64.67
70.62
64.97
72.25
75.18
74.63
68.26
66.37
72.94
66.38
138
Ba
65.29
68.09
69.02
71.71
68.09
66.79
68.17
66.78
67.71
71.06
74.02
73.28
68.22
65.99
74.83
71.23
68.67
66.49
67.76
65.35
70.28
68.00
66.33
71.24
69.14
71.79
67.54
70.16
65.53
68.71
65.72
68.60
70.58
65.86
67.28
70.23
68.36
67.25
67.62
9.52
70.77
75.91
73.07
71.27
69.94
65.49
70.97
69.22
71.06
74.02
73.28
68.22
65.99
74.83
71.23
175 Table D-1, continued.
Group
Sharpe 1
Sharpe 2
Sharpe 3
Sharpe 4
Sharpe 5
FrCase 1
FrCase 2
FrCase 3
Standard Ablation #
GSD
1
2
3
4
5
MACS-3
1
2
GSD
1
2
3
4
5
MACS-3
1
2
GSD
1
2
3
4
5
MACS-3
1
2
GSD
1
2
3
4
5
MACS-3
1
2
GSD
1
2
3
4
5
MACS-3
1
2
3
4
GSD
1
2
3
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
23
Na
26262.77
27044.68
27014.28
25880.31
26603.40
5430.07
5184.32
25867.00
26410.14
25574.98
25279.74
26627.21
5486.28
5128.97
29775.49
27945.15
28794.07
26316.05
24996.04
5268.58
5252.80
28662.37
28436.65
28081.59
27305.92
26437.96
5723.34
5350.85
25176.92
28697.86
27860.44
25222.54
27729.31
5579.69
5646.91
5470.06
5324.54
25512.68
27893.27
28327.32
5597.69
5146.10
24973.33
27056.86
26735.92
24747.38
13066.39
6179.24
26113.21
25598.42
26321.89
27805.50
5013.06
5225.25
24
Mg
21995.92
21691.28
21849.38
21403.03
21406.18
1741.23
1674.68
21073.41
20401.63
20304.71
20590.48
20997.41
1695.08
1565.78
25779.77
23315.00
22922.32
21288.62
20211.07
1631.99
1600.67
26117.91
25358.93
24480.55
24032.51
23554.40
1878.24
1755.44
21327.23
23506.28
21700.15
20767.22
21745.12
1677.49
1723.41
1676.61
1680.61
20841.23
21098.09
22425.40
1706.63
1612.82
21482.28
22078.71
21675.27
20420.63
3465.26
1973.59
22490.17
21055.19
21182.49
22091.68
1580.43
1641.74
43
Ca
50600.01
50600.00
50600.00
50600.00
50600.00
376933.03
376933.03
50599.99
50600.00
50599.99
50599.99
50600.00
376932.97
376932.94
50600.00
50599.99
50600.00
50600.00
50599.99
376932.94
376932.97
50599.99
50600.00
50600.00
50600.00
50599.99
376932.94
376932.94
50600.00
50600.00
50600.00
50600.00
50600.00
376933.00
376933.00
376933.00
376933.03
50600.00
50600.00
50600.00
376933.00
376933.00
50599.99
50599.99
50600.00
50600.00
376932.97
376932.94
50600.00
50600.01
50600.00
50600.00
376933.03
376933.06
55
Mn
208.55
206.53
206.86
203.47
201.67
476.28
454.86
202.75
200.40
197.89
198.42
199.28
472.08
453.16
223.77
211.25
215.14
202.09
191.69
468.52
452.90
241.32
235.52
230.45
221.47
228.31
526.01
486.18
201.88
209.33
205.69
210.09
199.86
462.60
441.82
456.66
423.62
201.22
208.33
204.99
464.08
438.03
210.67
208.25
209.05
217.34
421.05
449.92
201.88
199.54
206.28
211.39
463.51
443.14
86
Sr
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
N/A
67.17
70.78
69.14
69.64
66.75
8052.24
8033.45
7696.87
7093.09
71.55
69.30
67.02
8632.51
7925.01
69.84
69.34
69.96
70.69
4689.77
8334.17
69.28
69.26
68.33
69.44
7828.82
7837.25
88
Sr
73.78
69.87
69.91
68.51
68.21
7573.56
7312.77
72.00
69.13
69.09
68.68
67.36
7675.25
7292.89
70.49
70.69
70.40
70.18
68.18
7494.23
7417.88
68.84
67.93
67.94
67.99
67.35
7786.71
7240.67
69.26
71.39
71.06
69.78
68.03
8197.29
8345.91
7470.86
6922.43
71.18
71.69
66.52
8828.53
8331.20
70.40
70.65
71.36
69.39
4674.43
8619.36
70.65
65.33
67.08
70.20
8138.45
8011.88
137
Ba
68.41
69.51
68.16
66.56
66.68
63.84
60.59
65.86
66.42
64.49
64.95
63.95
61.80
59.70
68.57
72.40
74.06
71.46
67.78
60.96
61.64
71.31
69.03
69.20
67.74
65.47
74.04
64.54
67.89
71.03
66.76
66.79
66.86
62.49
59.37
54.39
54.46
67.96
70.59
66.37
69.76
58.98
68.08
70.48
69.89
65.03
28.08
74.17
68.18
65.43
66.56
69.23
64.66
59.90
138
Ba
70.48
68.58
66.65
68.14
66.75
70.69
66.77
68.60
66.61
64.25
65.27
66.43
68.96
65.71
70.82
72.82
73.80
71.55
65.63
65.71
70.81
66.46
64.18
63.24
62.32
60.31
75.31
68.69
67.51
70.65
67.80
63.78
67.82
81.64
78.12
80.09
74.29
66.09
70.53
68.74
88.87
80.87
68.64
70.62
70.46
65.09
29.62
99.38
69.21
64.87
67.10
67.98
83.56
79.98
176 Table D-1, continued.
Group
FrCase 4
Lewis&Clark 1
Lewis&Clark 2
Lewis&Clark 3
Lewis&Clark 4
Lewis&Clark 5
Lewis&Clark 6
Standard Ablation #
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
GSD
1
2
3
4
MACS-3
1
2
23
Na
24033.64
28851.68
26309.80
27768.54
5331.81
5058.47
26074.50
29739.90
27044.16
28148.89
4953.10
5538.91
26698.41
26593.72
25449.03
26976.60
5177.22
5618.80
25829.64
26790.98
27895.33
27367.61
5482.62
5645.92
26119.66
24715.68
23862.68
23523.90
4933.57
5121.06
27039.55
30207.03
27383.34
30841.54
5397.45
5306.66
27913.77
29812.98
27357.80
27852.29
5164.40
5389.05
24
Mg
21726.63
23488.34
22898.48
22683.50
1680.48
1549.75
21566.27
22397.66
21954.03
21533.74
1531.85
1766.34
21286.69
21824.11
21585.56
22237.19
1619.54
1615.69
20875.71
22295.04
22759.15
22576.00
1683.96
1693.18
21204.30
21291.68
21191.60
20349.85
1631.75
1655.20
21455.25
22686.40
21665.12
22441.57
1677.84
1633.58
21645.85
22872.73
21456.36
22311.46
1610.27
1638.23
43
Ca
50599.99
50599.99
50599.99
50599.99
376932.97
376932.94
50600.00
50600.00
50600.00
50600.00
376933.00
376933.00
50600.00
50599.99
50599.99
50599.99
376932.97
376932.97
50600.00
50600.00
50600.00
50600.01
376933.03
376933.03
50600.01
50600.01
50600.01
50600.01
376933.06
376933.03
50600.00
50600.00
50600.00
50600.00
376933.00
376933.00
50600.01
50600.01
50600.01
50600.01
376933.06
376933.06
55
Mn
208.76
212.20
207.51
209.81
436.47
423.06
205.88
197.58
206.76
200.84
425.49
446.41
212.06
199.92
205.91
208.86
428.96
430.60
197.01
203.74
198.06
202.52
432.64
439.56
194.45
198.45
193.68
201.30
418.50
435.81
208.68
220.13
208.24
216.97
445.41
444.58
209.61
222.11
212.92
221.01
439.03
450.49
86
Sr
67.54
69.56
64.15
70.84
7701.46
7258.07
70.91
66.66
67.16
70.96
7135.72
7491.98
72.71
68.28
67.09
71.52
7108.95
7734.90
70.13
68.56
65.30
67.54
7819.29
7708.11
71.99
74.06
70.07
74.27
8261.84
8762.46
69.93
74.23
65.43
70.05
8534.93
8039.00
67.94
72.40
65.86
68.91
7969.52
7897.62
88
Sr
70.46
68.34
64.15
71.84
8092.64
7300.24
70.62
72.79
67.18
69.14
7445.52
7717.06
72.79
68.39
67.25
67.75
7067.42
7969.96
72.45
70.97
72.26
70.91
8218.25
8100.29
74.47
70.28
75.35
70.59
8599.83
8947.10
68.68
80.17
67.54
69.34
8707.51
8160.09
67.93
76.09
67.20
65.53
8504.82
8037.54
137
Ba
65.55
68.82
62.28
68.06
61.07
56.70
69.27
76.11
66.59
67.90
55.82
59.38
71.52
68.34
64.21
65.61
53.58
60.56
71.51
70.46
71.86
70.93
65.37
60.89
73.11
64.64
63.21
66.16
61.53
62.50
66.23
77.17
69.21
70.93
63.25
60.66
66.68
73.36
71.16
66.74
64.24
60.61
138
Ba
62.35
69.64
62.80
68.46
79.53
73.77
70.39
77.66
66.18
70.77
69.85
78.37
69.24
69.52
64.12
66.41
73.87
80.02
72.03
70.58
72.32
69.57
84.10
83.47
72.71
67.37
64.22
64.63
83.62
84.09
65.87
73.75
67.21
72.59
85.42
84.22
66.63
71.74
67.44
70.18
82.17
84.41
177 Table D-2. Mean, 2*standard deviation, reproducibility (i.e., relative standard deviation,
RSD), and precision (i.e., deviation from true value) of standard reference material trace
element concentrations used for age-0 walleye otolith ablations in October 2013.
Standard Element
GSD
MACS-3
Mean
2*SD
RSD
Na
26884.54588
6044.582066
22.4835
24
Mg
22037.80657
5174.774104
23.4813
43
Ca
50600.00088
0.012618685
2.5E-05
55
Mn
207.9276471
34.5703111
86
Sr
69.35318182
88
Sr
23
True
Precision
21100
-4.444580894
16.6261
220
5.487433155
4.703053482
6.78131
69.4
0.067461357
69.84326733
4.679277815
6.69968
69.4
-0.638713727
137
Ba
68.22107843
13.26734125
19.4476
67
-1.822505121
138
67
-1.204565408
1756
3.116851645
Ba
67.80705882
13.16963041
19.4222
23
Na
5345.734894
794.1990638
14.8567
24
Mg
1701.268085
205.7831336
12.0959
43
Ca
376933.0096
0.080212484
2.1E-05
55
Mn
460.7161702
60.78144562
13.1928
536
14.04549063
86
Sr
7863.35913
918.2679945
11.6778
6700
-17.36356911
88
Sr
7788.169149
996.0116446
12.7888
6700
-16.24133058
137
Ba
64.17765957
10.90611573
16.9936
58.7
-9.331617674
138
Ba
75.57553191
14.88213508
19.6917
58.7
-28.74877669
178 Table D-3. Standard reference materials for adult walleye otoliths ablated in October
2013. USGS synthetic glass standard GSE-1G was used as the calibration standard, and
two additional reference standards (GSD-G1 and MACS-3) were used as quality controls
for verification of instrument accuracy and precision. Each standard was ablated in three
to five locations after every 4-10 samples to adjust for possible instrument drift.
Group
Standard
Adult 1
GSD
MACS-3
Adult 2
GSD
MACS-3
Adult 3
GSD
MACS-3
Adult 4
GSD
MACS-3
Adult 5
GSD
MACS-3
Adult 6
GSD
MACS-3
Adult 7
GSD
MACS-3
Ablation #
23
Na
24
Mg
43
Ca
55
Mn
86
Sr
88
Sr
137
Ba
138
Ba
1
2
3
4
28363.5
26804.61
25858.47
27054.64
22222.53
21872.95
21154.12
22121.79
50600
50600
50600
50600
219.43
216.11
204.95
215.71
69.56
70.95
67.38
67.75
71.05
71.59
68.41
65.75
68.9
69.02
68.06
65.02
68.76
68.22
66.22
66.35
5
1
2
3
1
2
3
4
5
1
2
1
2
3
4
5
1
2
26363.16
4111
4333.13
4283.3
26819.28
26013.25
26469.65
25048.68
26039.41
4624.63
4430.35
26194.43
28079.28
26992.08
28325.24
27304.37
4163.83
4649.31
21847.88
1495.56
1501.62
1456.65
21719.22
21733.28
21659.43
20871.59
20827.17
1626.53
1496.51
22075.04
21983.88
21157.64
22197.27
22390.55
1435.06
1593.01
50599.99
376932.97
376932.97
376932.97
50600
50600
50600
50600
50600
376933
376933
50600
50600
50600
50600
50600
376933.03
376933
208.41
365.82
378.22
372.85
231.72
238.68
215.49
213.21
200.94
444.99
454.17
202.12
209.34
196.06
205.49
201.17
450.12
485.97
66.9
7532.15
7502.4
7813.75
72.77
71
71.35
66.07
67.37
8119.04
7688.65
70.24
69.09
68.04
71.39
71.68
7251.99
7581.84
67.5
7604.31
7520.59
7872.55
73.85
70
69.56
66.18
68.47
8110.18
7563.84
71.31
68.08
69.48
70.7
73.51
7509.56
7989.95
64.91
46.56
46.3
44.44
76.33
69.01
68.48
63.91
68.64
46.54
45.94
69.83
70.82
67.29
69.38
69.91
43.98
49.11
65.12
45.92
46.98
45.37
74.37
67.68
69.16
64.36
68.03
45.86
46.73
68.26
71.72
68.72
70.99
69.71
43.71
49
1
25652.25
21263.32
50600
193.63
70.77
69.93
69.82
66.62
2
25184.31
20896.27
50600
193.39
70.03
70.39
67.2
66.76
3
25928.84
21439.65
50600
190.5
71.44
71.57
68.01
66.67
4
25100.04
21383.06
50600
188.01
70.57
70.41
66.6
65.78
5
25940.43
21408.49
50600
191.29
68.48
67.05
65.41
67.44
1
4520.08
1515.19
376933
472.58
7838.31
7837.79
47.98
48.39
2
4141.59
1462.72
376933
434.63
7821.23
7966
48.86
48.89
1
26117.74
21446.2
50599.99
231.13
63.38
63.83
64.9
66.34
2
27649.03
21089.49
50599.99
228.42
68.67
68.4
69.08
68.12
3
27420.98
21513.37
50599.99
237.77
69.89
70.3
71.28
68.47
4
26995.76
21591.49
50600
231.8
69.82
65.79
64.66
66.03
5
29324.45
22413.17
50599.99
233.43
68.45
69.21
71.38
73.01
1
4206.75
1315.7
376932.97
434.34
7595.42
7539.11
47.12
46.62
2
3778.67
1377.45
376932.97
410.09
7415.32
6795.28
42.65
41.59
1
27283.2
22133.3
50600
204.03
69.31
71.84
71.3
70.57
2
3
4
5
1
2
1
2
3
4
5
1
2
27042.13
27623.99
27775.84
26532.57
4269.07
3862.39
27147
26714.64
26678.81
26767.72
26539.24
4496.42
3800.85
21966.05
22269.19
21976.71
21890.13
1528.1
1313.65
21825.16
21618.46
22485.25
22057.13
22039.83
1554.44
1308.29
50600
50600.01
50600
50600
376933.03
376933.03
50600
50599.99
50600
50599.99
50599.99
376932.94
376932.94
201.06
193.35
202
204.94
442.66
389.59
196.16
201.33
194.17
207.43
209.04
451.75
426.17
70.49
70.19
67.87
69.37
7765.21
7231.25
67.53
70.03
70.23
69.15
71.11
8368.51
7370.36
71.46
70.39
67.27
67.73
7745.09
7317.13
70.14
71.46
68.48
65.83
68.96
8392.04
7357.06
68.82
65.35
65.58
64.8
49.42
43.69
69.9
67.36
63.44
65.82
64.72
50.01
42.55
70.02
67.83
65.27
65.8
46.11
40.58
69.3
68.26
66.59
65.24
65.73
48.64
40.48
179 Table D-4. Mean, 2*standard deviation, reproducibility (i.e., relative standard deviation,
RSD), and precision (i.e., deviation from true value) of standard reference material trace
element concentrations used for adult otolith ablations in October 2013.
Standard
Element
GSD
23
Na
26775.69 1911.754
7.139889213
24
Mg
21729.72
915.229
4.21187758
43
Ca
50600
0.009456
1.86873E-05
55
Mn
208.906
29.23155
86
Sr
88
Sr
MACS-3
Mean
2*SD
RSD
True
Precision
21100
-2.984436
13.99267953
220
5.0427273
69.38057 3.736642
5.385717982
69.4
0.0279951
69.31086 4.550996
6.566064637
69.4
0.1284479
137
Ba
67.85543 5.492526
8.094453535
67
-1.2767591
138
Ba
67.92914 4.578051
6.739450394
67
-1.3867804
1756
16.550949
23
Na
4244.758
557.663
13.13768604
24
Mg
1465.365 199.0234
13.58183013
43
Ca
55
86
88
0.059137
1.56889E-05
Mn
427.5967 73.40062
17.16585456
536
20.224502
Sr
7659.695 621.4137
8.112773648
6700
-14.323811
Sr
7674.699 764.9345
9.96696505
6700
-14.547741
46.34333 4.919603
10.61555742
58.7
21.050539
12.56161006
58.7
22.218058
137
Ba
138
Ba
376933
45.658
5.73538
180 Table D-5. Standard reference materials for adult walleye otoliths ablated in July 2014.
USGS synthetic glass standard GSE-1G was used as the calibration standard, and two
additional reference standards (GSD-G1 and MACS-3) were used as quality controls for
verification of instrument accuracy and precision. Each standard was ablated in three to
five locations after every 4-10 samples to adjust for possible instrument drift.
Site
Four Mile
Garrison Tailrace
64.6
62.76
65.79
73.35
70.12
74.01
66.37
64.91
61.72
71.54
68.72
71.69
4487.02 1990.18 376933
625.77 6201.08 7106.91
70.07
68.65
4824.13 2054.52 376933
644.81 5928.22 6804.99
67.35
70.14
3
4371.6
627.23
6009.9
7170.94
68
67.42
1
17074.4 20615.3
50600
260.73
60.92
76.51
67.6
68.16
2
17757.4 20808.2
50600
276.99
70
80.64
71.07
69.55
50600
273.36
77.69
78.51
1
2
GSD
GSD
GSD
GSD
GSD
GSD
MACS-3
Stanton
555.42 6683.4 7726.41
543.02 6946.79 7407.84
554.45 6800.58 7454.86
281.67 66.12
81.56
279.68
67.8
79.99
283.16
69.7
81.89
2015.54 376933
1989.62 376933
1916.93 376933
21011.8 50600
20698 50600
21164.1 50600
MACS-3
Porcupine
Ba
83.32
72.12
65
4244.68
4427.39
4081.68
18309.5
18312.9
19014.8
MACS-3
Peoria
137
1
2
3
1
2
3
MACS-3
Langliers
Ba
83.55
71.57
63.81
GSD
MACS-3
Na
Mg
20634.4 24486
18940.9 21826
17396 20360.1
43
MACS-3
MACS-3
Kneifel
24
Ablation #
1
2
3
MACS-3
Kimball Bottoms
23
Standard
GSD
Ca
50600
50600
50600
2051.18 376933
55
Mn
248.37
231.27
226.24
86
Sr
62.63
67.6
64.76
88
Sr
74.24
72.03
69.36
138
3
18090.8 20810.5
68.58
68.56
1
4892.59 2127.36 376933
627.05 6807.35 8226.04
71.45
72.96
2
4853.31 2159.67 376933
636.07
7561.14
70.87
69.34
3
4851.49 2116.76 376933
625.19 6629.12 7890.36
72.01
72.86
1
18273.5 21045.1
50600
279.78
64.82
81.82
70.45
72.14
2
17953.1 20972.7
50600
282.71
71.24
77.77
69.1
69.01
3
19149.8 21504.4
50600
286.18
68.18
79.04
68.52
68.55
1
4685.77 2085.92 376933
593.13 6491.47 8239.73
74.08
75.33
2
5371.03 2389.89 376933
666.17 6940.38 8508.11
81.68
82.65
3
5247.13
2145.5
376933
629.89 6581.08 8090.55
78.76
76.03
1
2
1
2
3
1
2
3
1
2
3
1
2
3
1
2
3
1
2
3
1
2
3
18684.2
18703.1
3924.04
4424.66
4562.78
22198.1
23565.3
22909.9
4678.47
4407.03
4654.35
17303.9
17220.5
17734.8
3806.7
4438.81
5167.73
20614.2
20389.5
19370.6
4316.36
4223
4544.07
20944.8
21770
1815.36
2036.1
2012.8
21589.9
21019.7
22240
1969.17
1928.65
1913
20723.4
20616.4
20630.9
1806.14
2071.76
2333.34
25236.5
23809.7
22289
1968.91
1906.23
1992.82
50600
50600
376933
376933
376933
50600
50600
50600
376933
376933
376933
50600
50600
50600
376933
376933
376933
50600
50600
50600
376933
376933
376933
240.99
247.49
528.01
536.99
619.42
1232.33
1312.08
1301.11
1216.33
1224.62
1187.49
253.3
251.85
262.18
543.47
606.17
605.93
274.53
244.99
247.48
554.59
531.31
557.32
72.49
70.42
61.3
71.19
74.05
2463.14
2621.51
2555.27
380.62
374.29
370.66
83.72
69.88
67.79
64.88
67.1
72.86
82.03
80.57
74.56
66.4
66.74
62.71
72.27
70.26
59.94
67.25
68.44
2680.53
2740.31
2819.7
425.49
378.17
410.27
81.43
69.48
68.26
64.3
73.21
73.49
82.94
79.36
75.58
65.52
64.46
64.39
6359.9
68.39
64.35
6493.48
6935.16
7165.31
2471.2
2551.28
2527.51
38083.1
37281.4
34543.3
70.17
67.93
68.85
6164.7
6601.65
7097
77.18
63.52
66.44
6933.84
6705.19
6878.32
74.21
71.08
7465.87
7453.49
7921.99
2591.1
2626.75
2609.72
41705.1
41299.4
38082.2
82.61
77.19
77.09
7325.58
7460.53
8110.82
76.84
72.94
74.46
7685.38
7479.48
7348.1
181 Table D-5, continued.
Site
Steckel
Standard
GSD
MACS-3
Vanderwall
GSD
MACS-3
Washburn
GSD
MACS-3
Ablation #
1
2
3
1
2
3
1
2
3
1
2
3
1
2
3
1
2
3
23
Na
18171.7
18228.6
18016.1
5336.62
5424.97
5166.24
19374.5
19138.8
18807.3
4376.97
4580.2
4279.07
20472.2
21036
19872.6
3807.03
4175.6
4334.84
24
Mg
21426.5
21559.2
20950.4
2402.43
2155.06
2167.71
22433.3
21963.5
21451.9
1894.98
1933.76
2007.32
24265.4
24792.5
22852.1
1883.12
1872.62
1867.93
43
Ca
50600
50600
50600
376933
376933
376933
50600
50600
50600
376933
376933
376933
50600
50600
50600
376933
376933
376933
55
Mn
270.43
273.01
273.21
648.48
650.89
627.93
225.37
241.39
244.83
530.78
556.6
565.01
258.24
250.15
237.67
532.44
552.04
514.13
86
Sr
62.24
65.3
65.75
6757.36
6038.19
6406.01
62.05
68.17
69.83
6903.16
6953.15
6899.52
81.88
64.92
66.08
6657.09
6549
7013.16
88
Sr
72.07
77.97
76.78
7808.83
7210.08
7485.79
69.8
71.12
72.96
7689.58
6984.28
7511.12
72.33
72.58
70.93
6961.77
7328.17
7824.7
137
Ba
65.48
67.82
63.98
74.05
67.67
71.06
73.88
70.23
69.18
68.61
62.04
67.92
70.5
79.04
78.7
60.34
61.97
64.59
138
Ba
67.87
70.59
69.18
76.16
71.76
73.26
74.11
71.85
69.32
65.96
61.27
63.31
72.57
80.81
78.1
58.54
61.63
62.56
182 Table D-6. Mean, 2*standard deviation, reproducibility (i.e., relative standard deviation,
RSD), and precision (i.e., deviation from true value) of standard reference material trace
element concentrations used for adult otolith ablations in July 2014.
Standard
Element
Mean
2*SD
RSD
GSD
23
Na
18760.20448
2212.943275
11.79594432
24
Mg
21828.19828
2792.04953
12.79102148
43
Ca
50599.99621
0.021655612
4.27977E-05
55
Mn
258.8706897
36.97659373
14.28380856
220
-17.6684953
86
Sr
67.74172414
9.475639547
13.9878925
69.4
2.389446487
88
Sr
75.73517241
7.975182256
10.53035466
69.4
-9.12849051
MACS-3
True
Difference
21100
-3.451176663
137
Ba
72.13793103
10.97328103
15.21152724
67
-7.668553783
138
Ba
72.49448276
9.909694512
13.66958441
67
-8.200720535
23
Na
4574.250333
911.3970438
19.92451172
24
Mg
2039.048667
306.0865986
15.01124537
1756
-16.11894457
43
Ca
376933.027
0.157318039
4.17363E-05
55
Mn
586.3236667
92.74059164
15.81730312
536
-9.388743781
86
Sr
6651.018667
666.244265
10.01717629
6700
0.731064677
88
Sr
-13.05643781
7574.781333
812.4894279
10.72624268
6700
137
Ba
68.43
10.27734032
15.01876417
58.7
-16.5758092
138
Ba
68.12766667
11.43047069
16.77801582
58.7
-16.06076093
183 Appendix E. Missouri River water trace elemental signatures (μmol/mol) based on
samples collected at sites throughout lakes Oahe, Sharpe, Francis Case, and Lewis &
Clark in summer 2012. Locations were not available for all sites. Samples were also
collected in winter 2014 and fall 2013. Fall 2013 samples were removed from analyses
because a flood altered trace element concentrations. However, water signatures were
temporally consistent between summer 2012 and winter 2014 for Sr:Ca (F57,1 = 69.89, P
= 0.10) and Ba:Ca (F57,1 = 102.62, P = 0.49), the elements used to investigate walleye
environmental history.
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Site
Knife R.
Knife R.
Heart R.
Heart R.
Hazelton
Hazelton
Cannonball R.
Cannonball R.
Beaver Creek
Beaver Creek
Beaver Bay
Beaver Bay
Fort Yates
Fort Yates
W. Pollock Emb.
W. Pollock Emb.
W. Pollock Main Channel
W. Pollock Main Channel
Grand R. Upstream
Mobridge Main Channel
Mobridge Main Channel
Moreau River Upstream
Swan Creek Emb.
Swan Creek Emb.
Swan Creek Main Channel
Swan Creek Main Channel
W. Whitlocks Emb.
W. Whitlocks Emb.
W. Whitlocks Main Channel
W. Whitlocks Main Channel
Bush's Main Channel
Bush's Main Channel
Cheyenne R. Upstream
Minneconjou Emb.
Minneconjou Emb.
Minneconjou Main Channel
Minneconjou Main Channel
Okobojo Emb.
Okobojo Emb.
Okobojo Main Channel
Okobojo Main Channel
Oahe Dam Face
Oahe Dam Face
Tailrace
Stilling Basin
Bad R. Upstream
Bad R. Upstream
Bad R. Confluence
Bad R. Confluence
Location
Location
Replicate
Na:Ca
Mg:Ca
N 47°19.362' W 101°22.544'
1A
6429.317 1205.996
N 47°19.362' W 101°22.544'
1B
6205.336 1173.927
1A
7632.587 1607.697
1B
7691.379 1604.199
N 46°31.645' W 100°32.558'
1A
2478.669 787.486
N 46°31.645' W 100°32.558'
1B
2513.152 793.754
1A
11508.940 2224.924
1B
11268.525 2172.645
N 46°15.706' W 100°23.977'
1A
3105.550 893.275
N 46°15.706' W 100°23.977'
1B
2997.096 886.509
N 46°14.514' W 100°31.231'
1A
2747.769 859.027
N 46°14.514' W 100°31.231'
1B
2730.083 851.603
N 46°05.730' W 100°35.407'
1A
2438.818 777.399
N 46°05.730' W 100°35.407'
1B
2393.682 767.419
N 44°30.081' W 100°38.606'
1A
2618.801 799.467
N 44°30.081' W 100°38.606'
1B
2604.254 799.565
N 45°33.000' W 100°20.629'
1A
2583.178 798.012
N 45°33.000' W 100°20.629'
1B
2597.279 793.595
1A
18586.037 1765.560
N 45°30.771' W 100°24.171'
1A
2686.020 799.800
N 45°30.771' W 100°24.171'
1B
2665.139 803.647
1A
8920.681 984.277
N 45°18.882' W 100°16.187'
1A
2769.459 806.593
N 45°18.882' W 100°16.187'
1B
2829.527 826.510
N 45°18.556' W 100°18.109'
1A
2732.540 805.913
N 45°18.556' W 100°18.109'
1B
2749.544 807.107
N 45°02.895' W 100°15.729'
1A
2723.542 785.361
N 45°02.895' W 100°15.729'
1B
2901.506 841.974
N 45°00.812' W 100°17.334'
1A
2699.367 792.998
N 45°00.812' W 100°17.334'
1B
2707.314 799.033
N 44°45.599' W 100°31.313'
1A
2702.201 781.487
N 44°45.599' W 100°31.313'
1B
2699.582 776.640
1A
1533.224 730.161
N 44°44.312' W 100°55.558'
1A
2665.070 755.916
N 44°44.312' W 100°55.558'
1B
2651.991 759.455
N 44°44.868' W 100°54.888'
1A
2661.660 759.642
N 44°44.868' W 100°54.888'
1B
2651.985 754.324
N 44°33.422' W 100°28.311'
1A
2630.791 767.724
N 44°33.422' W 100°28.311'
1B
2686.025 766.784
N 45°21.831' W 100°30.107'
1A
2745.560 787.926
N 45°21.831' W 100°30.107'
1B
2737.129 775.489
N 44°24.321' W 100°23.798'
1A
2762.925 773.932
N 44°24.321' W 100°23.798'
1B
2698.712 758.632
1A
2752.326 769.311
1A
2784.393 785.019
1A
5195.379 707.550
1B
5029.554 686.453
1A
2813.316 779.505
1B
2884.047 792.793
Mn:Ca
0.755
0.693
0.204
0.206
0.361
0.312
0.166
0.166
4.626
4.461
0.590
0.537
0.063
0.063
0.102
0.103
0.062
0.038
1.295
0.017
0.013
3.470
0.073
0.073
0.029
0.029
0.025
0.039
0.007
0.006
0.009
0.010
0.004
0.029
0.031
0.010
0.009
0.154
0.094
0.010
0.011
0.008
0.008
0.007
0.006
4.885
4.682
0.107
0.123
Sr:Ca
7.872
7.831
7.464
7.362
4.944
4.945
8.146
7.908
3.158
3.121
4.569
4.569
4.890
4.856
4.906
4.914
4.869
4.968
6.360
4.914
4.913
6.632
4.890
4.876
4.885
4.904
4.859
4.880
4.906
4.899
4.878
4.862
6.578
5.234
5.227
5.177
5.191
4.847
4.860
4.905
4.928
4.898
4.977
4.907
4.897
5.549
5.547
4.925
4.905
Ba:Ca
0.302
0.290
0.241
0.240
0.318
0.317
0.323
0.319
0.242
0.235
0.216
0.211
0.311
0.304
0.343
0.342
0.333
0.336
0.364
0.327
0.323
0.137
0.321
0.326
0.321
0.323
0.300
0.318
0.307
0.299
0.282
0.279
0.060
0.220
0.220
0.231
0.228
0.261
0.265
0.254
0.259
0.258
0.257
0.250
0.248
0.035
0.035
0.267
0.269
184 Appendix E, continued.
Reservoir
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Site
Hipple Lake
Antelope Creek
Antelope Creek
Degrey Emb.
Degrey Emb.
Degrey Main Channel
Degrey Main Channel
Joe Creek
Joe Creek
Iron Nations Emb.
Iron Nations Emb.
West Bend
West Bend
North Shore
North Shore
Tailrace
Tailrace
Stilling Basin
Stilling Basin
Cedar Shore
Cedar Shore
American Creek
American Creek
White River Upstream
White River Upstream
White River Confluence
White River Confluence
Elm Creek
Elm Creek
Platte Creek Emb.
Platte Creek Emb.
Platte Creek Main Channel
Platte Creek Main Channel
North Bay Emb.
North Bay Emb.
North Bay Main Channel
North Bay Main Channel
Tailrace Main Channel
Tailrace Main Channel
Tailrace Emb.
Tailrace Emb.
Niobrara R. Upstream
Niobrara R. Upstream
Niobrara R. Confluence
Niobrara R. Confluence
Running Water Main Channel
Running Water Main Channel
Running Water Side Channel
Running Water Side Channel
Location
Location
N 4903141
N 4903141
N 4902133
N 4902133
N 4887182
N 4887182
N 4883084
N 4883084
N 4891246
N 4891246
N 4879299
N 4879299
E 14 426155
E 14 426155
E 14 426100
E 14 426100
E 14 437061
E 14 437061
E 14 442307
E 14 442307
E 14 444158
E 14 444158
E 14 459023
E 14 459023
N 4841406
N 4841406
N 4839124
N 4839124
N 4822817
N 4822817
N 4795048
N 4795048
N 4792775
N 4792775
E 14 464093
E 14 464093
E 14 464508
E 14 464508
E 14 474426
E 14 474426
E 14 500270
E 14 500270
E 14 498808
E 14 498808
N 4767636
N 4767636
N 4766992
N 4766992
E 14 536632
E 14 536632
E 14 536283
E 14 536283
N 4735740
N 4735740
N 4735291
N 4735291
E 14 583438
E 14 583438
E 14 585845
E 14 585845
Replicate
1A
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
Na:Ca
3345.682
3237.168
3084.501
2549.360
2820.109
2753.030
2816.280
2819.875
2836.780
2799.006
2866.915
2721.626
2862.959
2811.306
2749.561
2687.994
2768.484
2718.619
2762.380
2781.946
2715.417
2009.855
1875.931
7008.372
7086.256
6967.926
7092.460
2800.512
2659.019
2723.473
2740.590
2722.291
2736.701
2696.259
2745.669
2789.186
2721.263
2834.907
2562.877
2615.834
2635.342
564.403
553.244
580.501
580.574
2675.476
2602.638
2617.870
2515.686
Mg:Ca
855.334
755.121
739.062
799.305
817.413
768.019
786.885
788.531
786.672
775.755
798.048
765.100
799.184
790.476
780.642
760.777
771.702
785.528
797.056
782.727
771.429
642.823
629.528
276.252
274.801
256.295
260.397
777.325
735.721
763.052
759.010
758.608
766.369
736.723
751.688
747.832
739.577
777.620
700.695
721.709
724.651
266.732
261.872
258.327
256.208
733.408
720.702
727.372
712.417
Mn:Ca
1.337
0.313
0.309
0.404
1.023
0.147
0.084
0.039
0.040
0.059
0.119
0.011
0.011
0.004
0.006
0.120
0.229
0.011
0.010
0.091
0.127
0.488
0.593
0.876
0.877
0.367
0.438
0.010
0.008
0.020
0.025
0.023
0.022
0.042
0.038
0.020
0.022
0.179
0.162
0.195
0.194
0.230
0.209
0.050
0.057
0.036
0.051
0.207
0.215
Sr:Ca
5.332
4.768
4.791
4.566
4.704
4.846
4.988
4.958
4.929
4.914
4.929
4.981
5.030
4.977
4.924
4.962
4.963
4.985
5.015
4.979
4.953
6.562
6.895
3.981
3.978
3.980
3.983
4.952
4.942
4.969
4.960
4.878
4.920
4.886
4.890
4.910
4.884
4.920
4.896
4.941
4.919
3.015
2.982
3.000
2.990
4.911
4.883
4.798
4.774
Ba:Ca
0.379
0.169
0.170
0.250
0.257
0.270
0.276
0.269
0.267
0.237
0.244
0.259
0.269
0.274
0.261
0.267
0.265
0.246
0.243
0.265
0.262
0.193
0.177
0.317
0.322
0.314
0.319
0.256
0.240
0.221
0.222
0.225
0.225
0.195
0.192
0.201
0.197
0.220
0.198
0.202
0.203
0.808
0.789
0.770
0.766
0.209
0.205
0.268
0.261
185 Appendix E, continued.
Reservoir
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Site
Springfield Backwater
Springfield Backwater
Springfield Main Channel
Springfield Main Channel
Springfield Emb.
Springfield Emb.
Tabor Main Channel
Tabor Main Channel
GPD
Location
N 4744489
N 4744489
N 4744489
N 4744489
N 4745728
N 4745728
Location
E 14 590470
E 14 590470
E 14 590470
E 14 590470
E 14 591219
E 14 591219
N 4745830
E 14 621846
Replicate
1A
1B
1A
1B
1A
1B
1A
1B
1A
Na:Ca
2818.925
2759.429
2709.141
2729.218
2261.898
2191.355
2766.231
2709.494
2674.834
Mg:Ca
740.237
725.017
719.783
725.238
667.376
648.410
732.265
731.377
710.214
Mn:Ca
0.135
0.167
0.060
0.062
0.566
0.621
0.010
0.011
0.023
Sr:Ca
5.111
4.992
4.883
4.915
4.452
4.342
4.903
4.819
4.876
Ba:Ca
0.447
0.429
0.223
0.227
0.193
0.191
0.243
0.241
0.235
186 Appendix F. Missouri River water trace elemental signatures (μmol/mol) based on
samples collected at sites throughout lakes Oahe, Sharpe, Francis Case, and Lewis &
Clark in winter 2014. Locations were not available for all sites. Samples were also
collected in winter 2014 and fall 2013. Fall 2013 samples were removed from analyses
because a flood altered trace element concentrations. However, water signatures were
temporally consistent between summer 2012 and winter 2014 for Sr:Ca (F57,1 = 69.89, P
= 0.10) and Ba:Ca (F57,1 = 102.62, P = 0.49), the elements used to investigate walleye
environmental history.
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Site
Knife R.
Knife R.
Knife R.
Knife R.
Knife R.
Knife R.
Knife R.
Knife R.
Knife R.
Knife R.
Heart R.
Heart R.
Heart R.
Heart R.
Heart R.
Heart R.
Heart R.
Heart R.
Heart R.
Heart R.
Hazelton
Hazelton
Cannonball R.
Cannonball R.
Cannonball R.
Cannonball R.
Cannonball R.
Cannonball R.
Cannonball R.
Cannonball R.
Cannonball R.
Cannonball R.
Beaver Creek
Beaver Creek
Beaver Bay
Beaver Bay
Beaver Bay
Beaver Bay
W. Pollock Emb.
W. Pollock Emb.
Grand R. Upstream
Grand R. Upstream
Grand R. Upstream
Grand R. Upstream
Grand R. Confluence
Grand R. Confluence
Grand R. Confluence
Grand R. Confluence
Mobridge Main Channel
Mobridge Main Channel
Location
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 47°19.362'
N 46°31.645'
N 46°31.645'
N 46°15.706'
N 46°15.706'
N 46°14.514'
N 46°14.514'
N 46°14.514'
N 46°14.514'
N 44°30.081'
N 44°30.081'
N 45°30.771'
N 45°30.771'
Location
Replicate
W 101°22.544'
1A
W 101°22.544'
1B
W 101°22.544'
2A
W 101°22.544'
2B
W 101°22.544'
3A
W 101°22.544'
3B
W 101°22.544'
4A
W 101°22.544'
4B
W 101°22.544'
5A
W 101°22.544'
5B
1A
1B
2A
2B
3A
3B
4A
4B
5A
5B
W 100°32.558'
1A
W 100°32.558'
1B
1A
1B
2A
2B
3A
3B
4A
4B
5A
5B
W 100°23.977'
1A
W 100°23.977'
1B
W 100°31.231'
1A
W 100°31.231'
1B
W 100°31.231'
2A
W 100°31.231'
2B
W 100°38.606'
1A
W 100°38.606'
1B
1A
1B
2A
2B
1A
1B
2A
2B
W 100°24.171'
1A
W 100°24.171'
1B
Na:Ca
5153.585
5142.006
5144.324
5080.396
5144.534
5073.547
5147.299
5127.741
5161.690
5142.880
5056.884
4828.725
4875.016
4854.633
5107.577
4945.823
5054.183
4909.561
4918.678
4865.475
2564.199
2597.942
4943.599
4677.226
4834.797
4950.704
4484.772
4635.096
4722.756
4764.527
4726.236
4810.017
2611.598
2616.440
2663.566
2631.362
2619.104
2657.433
2531.328
2493.285
10610.097
10420.660
10572.699
10409.858
2596.876
2518.983
2567.243
2535.305
2558.979
2504.404
Mg:Ca
1054.751
1059.979
1061.457
1043.609
1052.150
1043.934
1056.355
1060.059
1054.729
1056.270
1383.667
1330.911
1328.263
1325.723
1313.668
1308.997
1316.147
1341.221
1312.749
1353.553
774.780
789.377
1576.801
1492.620
1540.818
1580.381
1496.859
1473.447
1512.130
1521.214
1506.538
1537.402
800.906
789.661
791.884
795.885
781.447
797.327
781.895
771.895
1345.523
1328.322
1352.725
1317.717
800.197
772.343
794.114
784.229
796.142
796.924
Mn:Ca
0.836
0.825
0.814
0.792
0.819
0.822
0.843
0.819
0.833
0.819
0.029
0.009
0.006
0.007
0.010
0.008
0.009
0.007
0.009
0.007
0.001
0.001
0.367
0.352
0.375
0.381
0.392
0.375
0.379
0.375
0.376
0.372
0.011
0.008
0.029
0.026
0.032
0.038
0.010
0.010
0.030
0.033
0.021
0.037
0.019
0.021
0.009
0.009
0.004
0.003
Sr:Ca
7.076
7.196
7.063
7.083
7.145
7.171
7.082
7.035
7.172
7.163
6.404
6.443
6.394
6.447
6.407
6.478
6.426
6.401
6.420
6.425
4.636
4.694
6.435
6.456
6.394
6.422
6.439
6.503
6.482
6.448
6.459
6.465
4.470
4.465
4.431
4.704
4.681
4.808
4.660
4.630
6.235
6.149
6.178
6.084
4.732
4.628
4.687
4.653
4.646
4.604
Ba:Ca
0.226
0.229
0.228
0.224
0.229
0.229
0.230
0.228
0.233
0.230
0.210
0.212
0.213
0.209
0.208
0.210
0.209
0.207
0.210
0.210
0.260
0.260
0.119
0.118
0.117
0.117
0.117
0.119
0.117
0.118
0.116
0.117
0.260
0.251
0.262
0.261
0.271
0.281
0.262
0.257
0.215
0.215
0.221
0.226
0.265
0.253
0.261
0.263
0.256
0.251
187 Appendix F, continued.
Reservoir
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Oahe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Site
Moreau River Upstream
Moreau River Upstream
Moreau River Upstream
Moreau River Upstream
Moreau River Confluence
Moreau River Confluence
Moreau River Confluence
Moreau River Confluence
Moreau River Confluence
Moreau River Confluence
W. Whitlocks Emb.
W. Whitlocks Emb.
W. Whitlocks Main Channel
W. Whitlocks Main Channel
Cheyenne River Upstream
Cheyenne River Upstream
Cheyenne River Upstream
Cheyenne River Upstream
Cheyenne River Confluence
Cheyenne River Confluence
Cheyenne River Confluence
Cheyenne River Confluence
Cheyenne River Confluence
Cheyenne River Confluence
Minneconjou Emb.
Minneconjou Emb.
Okobojo Emb.
Okobojo Emb.
Oahe Dam Face
Oahe Dam Face
Stilling Basin
Stilling Basin
Bad R. Upstream
Bad R. Upstream
Bad R. Upstream
Bad R. Upstream
Bad R. Confluence
Bad R. Confluence
Bad R. Confluence
Bad R. Confluence
Bad R. Confluence
Bad R. Confluence
Location
Location
N 45°02.895'
N 45°02.895'
N 45°00.812'
N 45°00.812'
W 100°15.729'
W 100°15.729'
W 100°17.334'
W 100°17.334'
N 44°44.312'
N 44°44.312'
N 44°33.422'
N 44°33.422'
N 44°24.321'
N 44°24.321'
W 100°55.558'
W 100°55.558'
W 100°28.311'
W 100°28.311'
W 100°23.798'
W 100°23.798'
Replicate
1A
1B
2A
2B
1A
1B
2A
2B
3A
3B
1A
1B
1A
1B
1A
1B
2A
2B
3A
3B
4A
4B
5A
5B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
2A
2B
1A
1B
2A
2B
3A
3B
Na:Ca
6930.765
6831.375
6818.237
6815.617
6993.464
6902.233
7000.349
6903.695
6959.901
7000.182
2512.460
2475.958
2560.677
2531.020
2962.988
2069.677
2568.796
2774.035
2176.535
2008.954
2143.674
1943.148
2538.290
2069.909
2453.504
2450.044
2413.031
2398.566
2395.277
2418.250
2414.876
2431.320
2783.144
2763.746
2746.817
2756.644
2629.294
2614.979
2622.731
2649.593
2613.420
2631.344
Mg:Ca
955.233
962.567
965.929
963.471
997.112
989.820
973.246
981.527
962.923
1008.139
761.378
750.921
766.311
759.763
902.764
742.624
818.870
889.069
786.560
720.646
743.765
693.226
810.662
740.644
735.982
727.439
726.464
710.947
713.244
716.433
742.246
728.427
789.988
782.949
784.834
783.955
791.533
791.565
792.679
799.129
787.716
793.987
Mn:Ca
0.043
0.030
0.034
0.030
0.043
0.035
0.027
0.032
0.032
0.028
0.001
-0.001
0.001
0.000
0.011
0.134
0.012
0.009
0.136
0.138
0.141
0.139
0.008
0.135
-0.002
-0.001
0.000
0.000
-0.001
-0.001
-0.002
-0.002
0.016
0.009
0.012
0.010
0.011
0.013
0.013
0.015
0.014
0.014
Sr:Ca
5.347
5.395
5.525
5.610
5.549
5.674
5.579
5.429
5.795
5.512
4.433
4.402
4.515
4.397
5.336
5.883
5.315
5.311
5.779
5.883
5.881
5.919
5.332
5.896
4.486
4.566
4.553
4.496
4.466
4.521
4.445
4.423
5.295
5.327
5.322
5.263
5.241
5.285
5.272
5.288
5.285
5.271
Ba:Ca
0.136
0.131
0.138
0.140
0.140
0.137
0.135
0.141
0.138
0.139
0.226
0.227
0.232
0.227
0.233
0.064
0.231
0.232
0.063
0.063
0.063
0.063
0.232
0.064
0.178
0.177
0.185
0.184
0.184
0.186
0.178
0.176
0.206
0.207
0.210
0.208
0.220
0.218
0.215
0.217
0.223
0.218
188 Appendix F, continued.
Reservoir
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Sharpe
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Site
LaFramboise
LaFramboise
LaFramboise
LaFramboise
LaFramboise
LaFramboise
LaFramboise
LaFramboise
LaFramboise
LaFramboise
Hipple Lake
Hipple Lake
Hipple Lake
Hipple Lake
Hipple Lake
Hipple Lake
Hipple Lake
Hipple Lake
Hipple Lake
Hipple Lake
Antelope Creek
Antelope Creek
Antelope Creek
Antelope Creek
Antelope Creek
Antelope Creek
Antelope Creek
Antelope Creek
Antelope Creek
Antelope Creek
Joe Creek
Joe Creek
West Bend
West Bend
North Shore
North Shore
Cedar Shore
Cedar Shore
American Creek
American Creek
White R. Upstream
White R. Upstream
White R. Upstream
White R. Upstream
White R. Confluence
White R. Confluence
White R. Confluence
White R. Confluence
White R. Confluence
White R. Confluence
N 4887182
N 4887182
N 4891246
N 4891246
N 4879299
N 4879299
E 14 437061
E 14 437061
E 14 444158
E 14 444158
E 14 459023
E 14 459023
N 4841406
N 4841406
N 4841406
N 4841406
N 4839124
N 4839124
N 4839124
N 4839124
N 4839124
N 4839124
E 14 464093
E 14 464093
E 14 464093
E 14 464093
E 14 464508
E 14 464508
E 14 464508
E 14 464508
E 14 464508
E 14 464508
Replicate
1A
1B
2A
2B
3A
3B
4A
4B
5A
5B
1A
1B
2A
2B
3A
3B
4A
4B
5A
5B
1A
1B
2A
2B
3A
3B
4A
4B
5A
5B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
2A
2B
1A
1B
2A
2B
3A
3B
Na:Ca
2586.02
2582.92
2579.38
2566.21
2604.65
2592.58
2578.27
2584.82
2566.81
2556.80
3046.75
3041.55
3022.04
3034.24
3041.31
3065.38
3044.88
3031.22
3085.21
3067.15
3726.98
3706.37
3566.68
3695.99
3733.25
3741.63
3732.93
3727.30
3737.51
3749.65
2496.09
2467.34
2417.21
2422.69
2417.00
2442.12
2409.51
2434.58
1007.91
1011.92
2176.04
2140.95
2134.35
2123.82
2150.97
2161.43
2174.32
2101.32
2126.86
2140.56
Mg:Ca
801.08
799.56
802.35
797.99
803.28
803.81
805.93
803.34
800.85
797.24
837.67
837.77
836.42
834.05
832.92
837.04
841.79
835.05
841.91
839.01
522.70
521.85
507.27
522.54
524.29
526.71
524.52
521.50
525.16
523.54
735.85
732.05
717.39
724.09
728.60
719.68
723.38
723.09
480.96
478.00
286.04
285.40
276.03
278.47
276.98
268.97
285.30
280.59
278.99
278.32
Mn:Ca
0.03
0.03
0.00
0.00
0.03
0.03
0.00
0.00
0.03
0.03
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.00
0.40
0.40
0.42
0.45
0.36
0.36
0.40
0.38
0.43
0.42
0.01
0.01
0.01
0.01
0.01
0.00
0.00
0.00
0.06
0.07
0.02
0.19
0.25
0.23
0.02
0.05
0.31
0.19
0.39
0.02
Sr:Ca
5.24
5.28
5.29
5.26
5.26
5.31
5.28
5.25
5.25
5.30
5.32
5.35
5.39
5.38
5.38
5.40
5.40
5.38
5.39
5.38
4.49
4.50
4.32
4.47
4.55
4.47
4.49
4.51
4.45
4.51
4.95
4.84
6.47
6.52
5.21
5.21
4.42
4.44
8.12
8.16
3.31
3.44
3.24
3.33
3.35
3.37
3.34
3.38
3.30
3.30
Ba:Ca
0.23
0.22
0.23
0.22
0.23
0.23
0.22
0.22
0.22
0.23
0.26
0.26
0.26
0.26
0.25
0.25
0.26
0.25
0.25
0.25
0.07
0.07
0.08
0.08
0.08
0.07
0.07
0.07
0.07
0.07
0.19
0.19
0.22
0.21
0.19
0.19
0.18
0.18
0.06
0.06
0.34
0.35
0.36
0.37
0.34
0.32
0.34
0.38
0.38
0.34
189 Appendix F, continued.
Reservoir
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Francis Case
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Lewis & Clark
Site
Platte Creek Emb.
Platte Creek Emb.
Platte Creek Main Channel
Platte Creek Main Channel
North Bay Emb.
North Bay Emb.
North Bay Main Channel
North Bay Main Channel
Tailrace Main Channel
Tailrace Main Channel
Tailrace Emb.
Tailrace Emb.
Niobrara R. Upstream
Niobrara R. Upstream
Niobrara R. Upstream
Niobrara R. Upstream
Niobrara R. Confluence
Niobrara R. Confluence
Niobrara R. Confluence
Niobrara R. Confluence
Niobrara R. Confluence
Niobrara R. Confluence
Springfield Main Channel
Springfield Main Channel
Springfield Emb.
Springfield Emb.
Gavins Point Dam
Gavins Point Dam
N 4795048
N 4795048
N 4792775
N 4792775
E 14 500270
E 14 500270
E 14 498808
E 14 498808
N 4767636
N 4767636
N 4766992
N 4766992
E 14 536632
E 14 536632
E 14 536283
E 14 536283
N 4744489
N 4744489
N 4745728
N 4745728
N 4745830
N 4745830
E 14 590470
E 14 590470
E 14 591219
E 14 591219
E 14 621846
E 14 621846
Replicate
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
1A
1B
2A
2B
1A
1B
2A
2B
3A
3B
1A
1B
1A
1B
1A
1B
Na:Ca
2383.94
2473.75
2444.51
2415.92
2410.70
2411.87
2417.56
2393.75
2748.15
2616.68
2750.20
2727.44
424.12
424.81
400.08
424.96
405.55
417.09
423.05
419.71
430.59
416.98
2435.23
2625.15
2158.39
2087.51
2402.35
2461.89
Mg:Ca
686.54
706.79
701.53
695.90
721.38
733.23
720.07
708.04
792.56
798.74
819.27
848.13
252.98
251.10
241.44
252.34
244.72
251.52
248.18
251.06
245.18
250.71
755.50
821.65
745.42
702.50
699.41
720.90
Mn:Ca
0.01
0.02
0.03
0.02
0.01
0.01
0.01
0.01
0.17
0.17
0.48
0.53
0.02
0.02
0.01
0.01
0.02
0.01
0.02
0.02
0.01
0.01
0.00
0.01
0.01
0.01
2.23
2.23
Sr:Ca
4.27
4.41
4.30
4.28
4.55
4.63
4.77
4.85
5.20
5.23
5.18
5.21
2.83
2.75
2.77
2.74
2.77
2.78
2.77
2.76
2.78
2.81
5.12
5.05
6.78
6.92
52.10
54.81
Ba:Ca
0.17
0.18
0.17
0.17
0.19
0.19
0.19
0.19
0.21
0.21
0.21
0.20
0.91
0.88
0.88
0.87
0.88
0.90
0.88
0.87
0.89
0.90
0.25
0.25
0.25
0.25
1.48
1.57
190 Appendix G. Figures depicting spatial trends in Missouri River water chemistry that were
not used in chapters (i.e., manuscripts) above but were nevertheless fundamental for
investigating the utility of otolith microchemistry as a tool to study walleye
environmental history in Missouri River reservoirs.
191 Figure G-1. Mean water Sr:Ca values at 24 sites in Missouri River reservoirs (Oahe,
Sharpe, Francis Case [FC], Lewis & Clark [LC]) where age-0 Walleye were collected for
trace element analysis and site-specific chemical “blueprinting.” The error bars represent
SEs. Means with the same letter are not significantly different (ANOVA with Tukey’s
HSD test on log10 transformed values; P < 0.05). Site codes are as follows: BVB =
Beaver Bay, CAR = Cannonball River, CHR = Cheyenne River, GRR = Grand River,
HTR = Heart River, KNR = Knife River, MIN = Minneconjou Embayment, MOR =
Moreau River, OKO = Okobojo Embayment, WPL = West Pollock Embayment, WWE =
West Whitlocks Embayment, BAR = Bad River, FTG = Fort George, NSH = North
Shore, SSB = Sharpe Stilling Basin, WEB = West Bend, AMC = American Creek, NBE
= North Bay Embayment, PCE = Platte Creek Embayment, WHR = White River, GPD =
Gavin’s Point Dam, LCE = Lewis & Clark Embayment, NBR = Niobrara River, and SPE
= Springfield Embayment.
192 Figure G-2. Mean water Ba:Ca values at 24 collection sites in Missouri River reservoirs
(Oahe, Sharpe, Francis Case [FC], Lewis & Clark [LC]) where age-0 Walleye were
obtained for trace element analysis and site-specific chemical “blueprinting.” The error
bars represent SEs. Means with the same letter are not significantly different (ANOVA
with Tukey’s HSD test on log10 transformed values; P < 0.05). Site codes are as follows:
BVB = Beaver Bay, CAR = Cannonball River, CHR = Cheyenne River, GRR = Grand
River, HTR = Heart River, KNR = Knife River, MIN = Minneconjou Embayment, MOR
= Moreau River, OKO = Okobojo Embayment, WPL = West Pollock Embayment, WWE
= West Whitlocks Embayment, BAR = Bad River, FTG = Fort George, NSH = North
Shore, SSB = Sharpe Stilling Basin, WEB = West Bend, AMC = American Creek, NBE
= North Bay Embayment, PCE = Platte Creek Embayment, WHR = White River, GPD =
Gavin’s Point Dam, LCE = Lewis & Clark Embayment, NBR = Niobrara River, and SPE
= Springfield Embayment.
193 Appendix H. Water-otolith regressions that were not used in chapters (i.e., manuscripts)
above but were nevertheless fundamental for investigating the utility of otolith
microchemistry as a tool to study walleye environmental history in Missouri River
reservoirs. Fish and water sampling occurred in summer and fall 2012. Y-axes depict
mean terminal otolith element:Ca signatures for age-0 walleye. Juveniles were used under
the notion that their site fidelity would foster site-specific otolith signatures necessary for
chemical “blueprinting” of the Missouri River, an assumption that does not necessarily
apply for mobile, migratory adults.
194 Figure H-1. Linear regression of mean terminal otolith Sr:Ca values of age-0 Walleye on
water Sr:Ca signatures at collection sites in Lake Oahe tributaries, South Dakota and
North Dakota, USA. Fish and water sampling occurred in summer and fall 2012. The
error bars represent SEs.
195 Figure H-2. Linear regression of mean terminal otolith Sr:Ca values of age-0 Walleye on
water Sr:Ca signatures at collection sites in Lake Oahe tributaries, South Dakota and
North Dakota, USA. Fish and water sampling occurred in summer and fall 2012. The
error bars represent SEs.
196 Figure H-3. Linear regression of mean terminal otolith Sr:Ca values of age-0 Walleye on
water Sr:Ca signatures at collection sites in Lake Oahe tributaries, South Dakota and
North Dakota, USA, excluding the Cannonball River, which exhibited otolith terminal
Ba:Ca signatures. Even without the Cannonball River, the regression is still positive and
proportional. Fish and water sampling occurred in summer and fall 2012. The error bars
represent SEs.
197 Appendix I. Linear discriminant function analysis for age-0 Walleye collected in Lake
Oahe, North Dakota and South Dakota, USA. The first canonical function (CAN 1)
explained 58.86% of the variation in the data and discriminated among sites based on
terminal otolith Sr:Ca signatures. The second canonical function (CAN 2) accounted for
remaining data dispersion and discriminated among tributaries based on terminal otolith
Ba:Ca signatures. Labeled squares denote group centroids for collection sites. All
element:Ca ratios were log10 transformed prior to analysis.
198 Appendix J. Otolith transect diagrams that were not used in chapters (i.e., manuscripts)
above but were nevertheless fundamental for investigating the utility of otolith
microchemistry as a tool to study walleye environmental history in Missouri River
reservoirs.
199 Figure J-1. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected in
Beaver Bay, Lake Oahe, North Dakota, USA in Summer 2013. Site assignments reflect bivariate
(i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black triangle) displayed relatively stable
element:Ca ratios indicating prolonged site residence, whereas others (e.g., black circle) exhibited
variable signatures suggesting intra-reservoir movement. All transects proceed from the core of
the otolith toward the edge.
200 Figure J-2. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected in
Okobojo Bay, Lake Oahe, South Dakota, USA in Summer 2013. Site assignments reflect
bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black circle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others (e.g., white
circle) exhibited variable signatures suggesting intra-reservoir movement. All transects proceed
from the core of the otolith toward the edge.
201 Figure J-3. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected in
West Whitlocks Bay, Lake Oahe, South Dakota, USA in Summer 2013. Site assignments reflect
bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., white circle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others (e.g., black
triangle) exhibited variable signatures suggesting intra-reservoir movement. All transects proceed
from the core of the otolith toward the edge
202 Figure J-4. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected in
Bush’s Landing, Lake Oahe, South Dakota, USA in Summer 2013. Site assignments reflect
bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., white triangle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others (e.g., black
triangle) exhibited variable signatures suggesting intra-reservoir movement. All transects proceed
from the core of the otolith toward the edge.
203 Figure J-5. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected at
Oahe Dam Face (ODF), Lake Oahe, South Dakota, USA in Summer 2013. Site assignments
reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. MIN denotes Minneconjou Embayment.
Some adults (e.g., black circle) displayed relatively stable element:Ca ratios indicating prolonged
site residence, whereas others (e.g., white circle) exhibited variable signatures suggesting intrareservoir movement. All transects proceed from the core of the otolith toward the edge
204 Figure J-6. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected in
Joe Creek Embayment, Lake Sharpe, South Dakota, USA in Summer 2013. Site assignments
reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black circle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others (e.g., white
circle) exhibited variable signatures suggesting intra-reservoir movement and entrainment
through mainstem dams. All transects proceed from the core of the otolith toward the edge.
205 Figure J-7. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected in
North Shore Embayment, Lake Sharpe, South Dakota, USA in Summer 2013. Site assignments
reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., white circle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others (e.g., black
triangle) exhibited variable signatures suggesting intra-reservoir movement and entrainment
through mainstem dams. All transects proceed from the core of the otolith toward the edge.
206 Figure J-8. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult walleye collected in
Snake Creek Embayment, Lake Francis Case, South Dakota, USA in Summer 2013. Site
assignments reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black
circle) displayed relatively stable element:Ca ratios indicating prolonged site residence, whereas
others (e.g., white circle) exhibited variable signatures suggesting intra-reservoir movement and
entrainment through mainstem dams. All transects proceed from the core of the otolith toward the
edge.
207 Figure J-9. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult Walleye collected in
North Bay Embayment, Lake Francis Case, South Dakota, USA in Summer 2013. Site
assignments reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black
triangle) displayed relatively stable element:Ca ratios indicating prolonged site residence,
whereas others (e.g., black circle) exhibited variable signatures suggesting intra-reservoir
movement and entrainment through mainstem dams. All transects proceed from the core of the
otolith toward the edge.
208 Figure J-10. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult Walleye collected in
Lewis & Clark Tailrace, Lewis & Clark Lake, South Dakota, USA in Summer 2013. Site
assignments reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., black
triangle) displayed relatively stable element:Ca ratios indicating prolonged site residence,
whereas others (e.g., black circle) exhibited variable signatures suggesting intra-reservoir
movement and entrainment through mainstem dams. All transects proceed from the core of the
otolith toward the edge.
209 Figure J-11. Representative (a) Sr:Ca and (b) Ba:Ca otolith trasects of adult Walleye collected at
Tabor Landing, Lewis & Clark Lake, South Dakota, USA in Summer 2013. Site assignments
reflect bivariate (i.e., Sr:Ca, Ba:Ca) otolith signatures. Some adults (e.g., white circle) displayed
relatively stable element:Ca ratios indicating prolonged site residence, whereas others (e.g., black
circle) exhibited variable signatures suggesting intra-reservoir movement and entrainment
through mainstem dams. All transects proceed from the core of the otolith toward the edge.
210 Appendix K. Tables used for quantitative assessment of walleye entrainment that were
not included in chapters (i.e., manuscripts) above but were nevertheless fundamental for
investigating the utility of otolith microchemistry as a tool to study walleye
environmental history in Missouri River reservoirs.
211 Table K-1. Entrainment of adult walleye into Missouri River reservoirs during the 2011
flood as a percentage of total entrained adult walleye across years.
Collection Reserovir
Oahe
Sharpe
FC
LC
% (#) Into Sharpe
-57.14 (4)
10.00 (1)
21.42 (3)
% (#) Into Francis Case
--60.00 (6)
21.43 (3)
% (#) Into Lewis & Clark
---29.00 (4)
212 Table K-2. Summary of adult Walleye entrainment during the 2011 flood by age class.
Reservoir
Sharpe
Sharpe
Sharpe
Sharpe
Francis Case
Francis Case
Francis Case
Lewis & Clark
Lewis & Clark
Lewis & Clark
Total/Overall
Site
# Age-3+ Entrained % Entrained (Age-3+)
Joe Creek
0
0.00
N. Shore
1
100.00
Stilling Basin
2
100.00
W. Bend
2
66.67
Chamberlain Area
2
50.00
N. Bay
1
100.00
Snake Creek
2
100.00
GPD
3
75.00
LC Tailrace
1
50.00
Tabor
1
50.00
15
68.18
# Age-2 Entrained % Entrained (Age-2)
1
100.00
0
0.00
0
0.00
1
33.33
2
50.00
0
0.00
0
0.00
1
25.00
1
50.00
1
50.00
7
31.82
213 Appendix L. Tables that were not used in chapters (i.e., manuscripts) above but were
nevertheless fundamental to assessing changes in aquatic habitat in the Lewis & Clark
Delta after the 2011 flood.
214 Table L-1. UTM northing and easting coordinates for equidistant transects used to
evaluate changes in aquatic habitat in the Lewis & Clark Delta after the 2011 flood.
Landsat landcover geodatabases representing pre-flood (June 2010) and post-flood
(October 2011) conditions were displayed in ARCMap 10.1, and transects were overlaid
on imagery to facilitate aquatic habitat analyses, including enumeration of channels,
backwaters, and sandbars and measurement of main channel width and largest-backwater
area.
Transect
1
2
3
4
5
6
7
8
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
Northing
576906.51
577491.15
578075.79
578660.43
579245.08
579829.72
580414.36
580999.00
581583.65
582168.29
582752.93
583337.57
583922.21
584506.86
585091.50
585676.14
586260.78
586845.42
587430.07
588014.71
588599.35
589183.99
589768.63
590353.28
590937.92
591522.56
592107.20
592691.85
593276.49
593861.13
594445.77
Easting
4734799.94 Niobrara River
4735091.03
4734800.94
4735673.20
4734801.94
4736255.38
4734802.94
4736837.55
4734803.94
4737419.72
4734804.94
4738001.89
4734805.94
4738584.07
4734806.94
4739166.24
4734807.94
4739748.41
4734808.94
4740330.58
4734809.94
4740912.76
4734810.94
4741494.93
4734811.94
4742077.10
4734812.94
4742659.28
4734813.94
4743241.45
4743532.53 Downstream Limit of Delta