MOVEMENT OF SCAPHIRHYNCHUS SPECIES IN THE MISSOURI RIVER ABOVE
FORT PECK RESERVOIR, MONTANA
by
Ryan Roy Richards
A thesis submitted in partial fulfillment
of the requirements for the degree
of
Master of Science
in
Fish and Wildlife Management
MONTANA STATE UNIVERSITY
Bozeman, Montana
November 2011
©COPYRIGHT
by
Ryan Roy Richards
2011
All Rights Reserved
ii
APPROVAL
of a thesis submitted by
Ryan Roy Richards
This thesis has been read by each member of the thesis committee and has been
found to be satisfactory regarding content, English usage, format, citation, bibliographic
style, and consistency and is ready for submission to The Graduate School.
Dr. Christopher S. Guy
Approved for the Department of Ecology
Dr. David W. Roberts
Approved for The Graduate School
Dr. Carl A. Fox
iii
STATEMENT OF PERMISSION TO USE
In presenting this thesis in partial fulfillment of the requirements for a master’s
degree at Montana State University, I agree that the Library shall make it available to
borrowers under rules of the Library.
If I have indicated my intention to copyright this thesis by including a copyright
notice page, copying is allowable only for scholarly purposes, consistent with “fair use”
as prescribed in the U.S. Copyright Law. Requests for permission for extended quotation
from or reproduction of this thesis in whole or in parts may be granted only by the
copyright holder.
Ryan Roy Richards
November 2011
iv
DEDICATION
I dedicate this thesis to my late grandfather James Cecil Brickey. Thank you for
wanting to see me catch that fish worse than I did, and for having the patience to bait my
hooks and untangle my line. Thank you for Froggie Went A-Courtin’ and for every
single day you took me outdoors as a child. But most of all, thank you for leading by
example and for teaching me at an early age that the world is like a ripe huckleberry at
my fingertips.
v
ACKNOWLEDGEMENTS
I thank Michael Michelson and Nick Smith for their help in the field; I truly
appreciate both of your efforts and will not forget the hot, buggy hours we spent together.
Thanks to Bill Gardner of MT Fish, Wildlife and Parks, without his extraordinary efforts,
expertise, and support this project would not have been possible. For teaching me the
practical aspects of navigating the Missouri River, trammel netting, performing fish
surgeries, and tracking fish, thanks to Casey Jensen, Eli McCord, Randy Rodencal, and
Mike Wente.
Thanks to the U.S. Bureau of Reclamation and PPL Montana for supporting and
funding this research, with a special thanks to Sue Camp and Steve Leathe. Thanks to
Matt Jaeger for providing code and support for analyses, and to Eli Cureton and Joel Van
Eenennaam for training and assistance with histological analyses. Much gratitude to my
fellow graduate students, with special thanks to the Bens, Mike Meeuwig, Justin Spinelli,
John Syslo, Peter Brown, and Mariah Talbott.
I am forever indebted to my advisor Dr. Christopher Guy for his personal
attention and patience throughout the course of my education, I have truly learned much
from you. Thank you to Dr. Molly Webb for her input on the design and implementation
of the study and her invaluable expertise in all things physiological. Special thanks to Dr.
Robert Bramblett for encouraging me in my academic endeavors through the years, and
for showing me that scientists can still rock and roll.
Last but certainly not least I must express my unending gratitude to my beautiful
wife Crystal Lynette. With you by my side I can achieve anything.
vi
TABLE OF CONTENTS
1. INTRODUCTION TO THESIS ................................................................................... 1
Overview of Thesis ...................................................................................................... 3
References ................................................................................................................... 6
2. MOVEMENTS OF SCAPHIRHYNCHUS SPP.
IN THE UPPER MISSOURI RIVER ........................................................................... 9
Contribution of Authors and Co-Authors ..................................................................... 9
Manuscript Information Page ..................................................................................... 10
Abstract ..................................................................................................................... 11
Introduction ............................................................................................................... 12
Methods..................................................................................................................... 15
Fish Capture and Surgery ................................................................................... 15
Tracking............................................................................................................. 16
Hydrographs ...................................................................................................... 17
Movement and Movement Rates ........................................................................ 18
Results ....................................................................................................................... 19
Hydrographs ...................................................................................................... 19
Movement and Movement Rates ........................................................................ 20
Discussion ................................................................................................................. 22
Acknowledgements.................................................................................................... 27
Tables ........................................................................................................................ 28
Figures....................................................................................................................... 33
References ................................................................................................................. 38
3. SPAWNING RELATED MIGRATION OF SHOVELNOSE
STURGEON IN THE MISSOURI RIVER ABOVE FORT PECK
RESERVOIR, MONTANA ....................................................................................... 43
Contribution of Authors and Co-Authors ................................................................... 43
Manuscript Information Page ..................................................................................... 44
Abstract ..................................................................................................................... 45
Introduction ............................................................................................................... 46
Study Area ................................................................................................................. 48
Methods..................................................................................................................... 49
Fish Capture and Surgery ................................................................................... 49
Reproductive Assessment................................................................................... 50
Hydrographs and Water Temperature Profiles .................................................... 51
Tracking............................................................................................................. 52
Movement and Movement Rates ........................................................................ 53
vii
TABLE OF CONTENTS – CONTINUED
Substrate ............................................................................................................ 54
Data Analysis ..................................................................................................... 54
Results ....................................................................................................................... 56
Hydrographs and Water Temperature Profiles .................................................... 56
Movement and Movement Rates ........................................................................ 57
Substrate ............................................................................................................ 62
Discussion ................................................................................................................. 63
Acknowledgements.................................................................................................... 69
Tables ........................................................................................................................ 70
Figures....................................................................................................................... 76
References ................................................................................................................. 89
4. CONCLUSIONS ....................................................................................................... 96
References ............................................................................................................... 104
REFERENCES ............................................................................................................ 107
APPENDIX A: Shovelnose Sturgeon Reproductive Physiology Data .......................... 115
viii
LIST OF TABLES
Table
Page
2.1. Sample sizes for juvenile pallid sturgeon, adult pallid sturgeon,
and shovelnose sturgeon in the Missouri River above
Fort Peck Reservoir 2006-2009 ............................................................................... 28
2.2. Minimum, maximum, and mean (SD) daily discharge (m3/s)
and daily discharge rate of change (ROC) (m3/s/d) for the Missouri River
upstream of Fort Peck Reservoir for all hydrograph phases, 2006- 2009 ................. 29
2.3. Minimum, maximum, and mean ± 90% confidence interval (CI)
net and total movement rates (km/d) for juvenile pallid sturgeon in
the Missouri river above Fort Peck Reservoir for all hydrograph
phases 2006-2009 ................................................................................................... 30
2.4. Minimum, maximum, and mean ± 90% confidence interval (CI)
net and total movement rates (km/d) for adult pallid sturgeon in
the Missouri river above Fort Peck Reservoir for all hydrograph
phases 2006-2009 ................................................................................................... 31
2.5. Minimum, maximum, and mean ± 90% confidence interval (CI)
net and total movement rates (km/d) for shovelnose sturgeon in
the Missouri river above Fort Peck Reservoir for all hydrograph
phases 2006-2009 ................................................................................................... 32
3.1. Sample size and tagging location of telemetered shovelnose sturgeon
in the Missouri River above Fort Peck Reservoir 2008-2009 ................................... 70
3.2. Minimum, maximum, and mean (SD) daily discharge, daily water temperature,
And rates of change (ROC) for the Missouri River above Fort Peck Reservoir
for all hydrograph phases 2008-2009 ...................................................................... 71
3.3. Minimum, maximum, and mean ± 90% confidence interval (CI)
net and total movement rates (km/d) for confirmed spawning female
shovelnose sturgeon in the Missouri river above Fort Peck Reservoir
for all hydrograph phases 2008-2009 ...................................................................... 72
3.4. Minimum, maximum, and mean ± 90% confidence interval (CI)
net and total movement rates (km/d) for reproductive categories
of shovelnose sturgeon in the Missouri river above Fort Peck Reservoir
for all hydrograph phases in 2008 ........................................................................... 73
ix
LIST OF TABLES – CONTINUED
Table
Page
3.5. Minimum, maximum, and mean ± 90% confidence interval (CI)
net and total movement rates (km/d) for reproductive categories
of shovelnose sturgeon in the Missouri river above Fort Peck Reservoir
for all hydrograph phases in 2009 ........................................................................... 74
3.6. Likelihood chi-square statistics for overall selection
of substrate types by shovelnose sturgeon reproductive category
in the Missouri River above Fort Peck Reservoir during the suitable
spawning phase in 2009 (α = 0.1)............................................................................ 75
x
LIST OF FIGURES
Figure
Page
2.1. Map of Missouri and Marias rivers in Montana, with study area in detail ................ 33
2.2. Hydrograph phases for the Missouri River upstream of
Fort Peck Reservoir, MT 2006-2009, as defined by mean daily discharge ............... 34
2.3. Plots of river kilometer-standardized movement patterns of
juvenile pallid sturgeon and adult pallid sturgeon in the Missouri River
above Fort Peck Reservoir, 2006-2009.................................................................... 35
2.4. Plots of river kilometer-standardized movement patterns of
shovelnose sturgeon in the Missouri River above Fort Peck Reservoir,
2006-2009............................................................................................................... 36
2.5. Maps of the Missouri River above Fort Peck Reservoir showing
the distributions of juvenile pallid sturgeon, adult pallid sturgeon,
and shovelnose sturgeon locations 2006-2009 ......................................................... 37
3.1. Map of Missouri and Marias rivers in Montana, with study area in detail ................ 76
3.2. Hydrograph phases for the Missouri River upstream of
Fort Peck Reservoir, 2008 and 2009 as defined by discharge
and water temperature relative to shovelnose sturgeon putative
reproductive life history .......................................................................................... 77
3.3. Weekly mean net movement rates of confirmed spawning female
shovelnose sturgeon in the Missouri River upstream of Fort Peck Reservoir
2008-2009 as a function of weekly mean discharge and water temperature ............. 78
3.4. Weekly mean total movement rates of confirmed spawning female
shovelnose sturgeon in the Missouri River upstream of Fort Peck Reservoir
2008-2009 as a function of weekly mean discharge and water temperature ............. 79
3.5. Mean net movement rates for male, confirmed spawning female,
potentially spawning female, and atretic female shovelnose sturgeon
by hydrograph phase in 2008 .................................................................................. 80
3.6. Mean total movement rates for male, confirmed spawning female,
potentially spawning female, and atretic female shovelnose sturgeon
by hydrograph phase in 2008 .................................................................................. 81
xi
LIST OF FIGURES – CONTINUED
Figure
Page
3.7. Mean net movement rates for male, confirmed spawning female,
potentially spawning female, atretic female, and reproductively inactive
female shovelnose sturgeon by hydrograph phase in 2009 ....................................... 82
3.8. Mean total movement rates for male, confirmed spawning female,
potentially spawning female, atretic female, and reproductively inactive
female shovelnose sturgeon by hydrograph phase in 2009 ....................................... 83
3.9. Movement patterns of male, confirmed spawning female,
potentially spawning female, and atretic female shovelnose sturgeon
in the Missouri River above Fort Peck Reservoir, 2008 ........................................... 84
3.10. Movement patterns of male, confirmed spawning female,
potentially spawning female, atretic, and reproductively inactive female
shovelnose sturgeon in the Missouri River above Fort Peck Reservoir, 2009 ......... 85
3.11. Mean river kilometer of confirmed spawning female shovelnose sturgeon
During suitable spawning conditions in the Missouri river above Fort Peck
Reservoir, 2008 and 2009...................................................................................... 86
3.12. Distribution of substrate types in the Missouri River upstream of
Fort Peck Reservoir, 2009, as estimated from 558 randomly chosen points ........... 87
3.13. Substrate use by shovelnose sturgeon reproductive category versus
availability in the Missouri River upstream of Fort Peck Reservoir
during the suitable spawning phase, 2009 .............................................................. 88
xii
ABSTRACT
Some Scaphirhynchus spp. populations are endangered, in decline or extirpated.
Operation of dams and reservoirs on the Missouri River has been implicated in declines
in Scaphirhynchus spp. abundance. It is hypothesized that discharge from dams upstream
of Scaphirhynchus spp. populations are insufficient to provide the environmental cue
initiating spawning migrations. The goal of this thesis was to investigate the effects of
variations in spring discharge on movements of Scaphirhynchus spp. in the Missouri
River above Fort Peck Reservoir. A secondary goal was to provide information on the
distribution of Scaphirhynchus spp. locations relative to larval drift distance. Twentyfour hatchery-reared juvenile pallid sturgeon, seven adult pallid sturgeon, and 192
shovelnose sturgeon were tracked from 2006 through 2009 and movements were
compared among years with hydrologic conditions from above to below average
discharge. Seventy-eight shovelnose sturgeon in five reproductive categories (i.e., males,
confirmed spawning females, potentially spawning females, atretic females, and
reproductively inactive females) were tracked in 2008 and 2009. Movements were
compared among all reproductive categories within years and between years for
confirmed spawning females while integrating the environmental effects of discharge and
water temperature. The majority of pallid sturgeon locations in all years were within 75
km of the headwaters of Fort Peck Reservoir. Shovelnose sturgeon locations were
distributed across the entire study reach, and were 100 km further upstream than pallid
sturgeon locations. Based on current estimates, an insufficient length of river exists
upstream of the Fort Peck Reservoir headwaters for pallid sturgeon larval drift.
Movement rates of Scaphirhynchus spp. did not differ among years 2006-2009, and
movements did not differ between years for confirmed spawning female shovelnose
sturgeon 2008-2009, indicating that discharge did not influence movements. At the
conditions in this study, movement rates of confirmed spawning female shovelnose
sturgeon were highest at water temperatures suitable for spawning regardless of
discharge, providing support for the hypothesis that water temperature rather than
discharge is a more likely proximate cue initiating spawning migrations in
Scaphirhynchus spp.
1
CHAPTER ONE
INTRODUCTION TO THESIS
Pallid sturgeon Scaphirhynchus albus and shovelnose sturgeon Scaphirhynchus
platorynchus (collectively referred to as Scaphirhynchus spp.) are native to the Missouri
and Mississippi River basins and are sympatric throughout much of their ranges (Bailey
and Cross 1954; Kallemeyn 1983; Keenlyne 1997). Currently the pallid sturgeon is listed
as endangered (Dryer and Sandvol 1993) and although generally stable, some shovelnose
sturgeon populations are in decline or extirpated (Koch and Quist 2010). In the Missouri
River downstream of Morony Dam near Great Falls, Montana, to the headwaters of Fort
Peck Reservoir (hereafter referred to as the upper Missouri River), shovelnose sturgeon
have continued to recruit. However, no natural pallid sturgeon recruitment has been
documented for at least 40 years, and estimates of the number of wild adult pallid
sturgeon in the upper Missouri River varied from 40 to 100 in 2003 (Webb et al. 2005).
Anthropogenic activities that alter riverine habitat likely have direct impacts on
Scaphirhynchus spp., and operation of dams and reservoirs on the Missouri River has
been implicated in declines in Scaphirhynchus spp. abundance (Hesse et al. 1989; Dryer
and Sandvol 1993; USFWS 2000, 2003; Galat et al. 2005; DeLonay et al. 2007).
Dams and reservoirs fragment fish habitat, alter and decouple natural discharge
and water temperature regimes, reduce sediment transport and disconnect the floodplain
from the river channel (Junk et al. 1989; Sparks et al. 1990; Stanford et al. 1996; Poff et
al. 1997). Several dams exist in the Missouri River basin upstream of the study area.
2
Three dams operated by the U.S. Bureau of Reclamation (USBR) have the most effect on
discharge and habitat, i.e, Canyon Ferry Dam on the Missouri River near Helena,
Montana (rkm 3,626, measuring from the confluence with the Mississippi River), Tiber
Dam on the Marias River near Chester, Montana (rkm 129, measuring from confluence
with the Missouri River), and Gibson Dam on the Sun River (rkm 163, measuring from
confluence with the Missouri River) (Gardner and Jensen 2011). Under Section 7 of the
Endangered Species Act, the USBR has initiated a study on water operations at its
facilities in the Missouri River basin above Fort Peck Reservoir. Of utmost concern is
the effect of these operations on Scaphirhynchus spp. populations. Rising discharge
associated with the spring pulse is hypothesized to cue upstream migrations in
Scaphirhynchus spp. (Becker 1983; Keenlyne and Jenkins 1993; Goodman 2009; Tripp et
al. 2009), and regulation has altered the spring discharge of the Missouri River (Galat et
al. 2001). Based on the hypothesis that rising spring discharge is a spawning cue for
pallid sturgeon, the U.S. Fish and Wildlife Service (USFWS) has required that dams
operated by the U.S. Army Corps of Engineers on the Missouri River be managed for
increased spring discharge (USFWS 2000, 2003). Previous operational discharge was
believed to be insufficient to initiate migrations in pallid sturgeon and other native fishes
(USFWS 2003). However, little research has been conducted investigating the effects of
variations in spring discharge on movements of Scaphirhynchus spp. Understanding how
movements differ under varying spring discharge scenarios will provide baseline data to
help guide future release schedules designed to benefit Scaphirhynchus spp. populations.
3
In addition to altering natural discharge regimes, dams affect the amount of
habitat available for multiple Scaphirhynchus spp. life-history needs (Braaten et al. 2008;
Gerrity et al. 2008). White sturgeon Acipenser transmontanus populations in fragmented
river reaches terminating in reservoirs have reduced recruitment (Jager et al. 2002), and it
has been hypothesized that an insufficient length of free-flowing river downstream from
spawning locations is a mechanism for the lack of recruitment in pallid sturgeon in the
upper Missouri River (Braaten et al. 2008). Thus, documenting movement patterns and
the distribution of Scaphirhynchus spp. locations relative to larval drift distance may
identify research priorities in the upper Missouri River.
Overview of Thesis
I examined the influence of variation in the annual spring pulse of the Missouri
River above Fort Peck Reservoir on movements of shovelnose sturgeon and pallid
sturgeon. I was particularly interested in comparing movements among years with
differing discharge rate of change, peak magnitude, duration, and timing because rising
discharge associated with the spring pulse is hypothesized to cue upstream migrations in
Scaphirhynchus spp. (Becker 1983; Keenlyne and Jenkins 1993; Goodman 2009; Tripp et
al. 2009). It has been hypothesized that the extended larval drift observed in pallid
sturgeon may be contributing to the lack of recruitment observed in the upper Missouri
River (Braaten et al. 2008); thus, movement patterns and the distributions of
Scaphirhynchus spp. locations relative to larval drift distance were also compared. Care
must be taken when interpreting the results of this study, as telemetry studies that try to
4
link movement and habitat selection to environmental variables such as discharge and
temperature are correlative in nature and do not necessarily imply cause and effect
(Rogers and White 2007).
In chapter two, based on the hypothesis that rising spring discharge is a cue for
migration in Scaphirhynchus spp., the upper Missouri River hydrographs (2006-2009)
were divided into three phases defined by discharge. Movement patterns and movement
rates of juvenile pallid sturgeon, adult pallid sturgeon, and shovelnose sturgeon were
compared among years and among periods of rising discharge (ascending limb),
decreasing discharge (descending limb), and baseflow (post-descending limb). The
specific objectives of chapter two were to (1) compare the movements of juvenile pallid
sturgeon, adult pallid sturgeon, and shovelnose sturgeon among years with differing
spring hydrographs, and (2) compare movement patterns and distributions of juvenile
pallid sturgeon, adult pallid sturgeon, and shovelnose sturgeon locations in the upper
Missouri River. In chapter two I hypothesized that movements of Scaphirhynchus spp.
would be greater in a year with higher discharge, the highest movement rates would be
coincident to rising discharge, movements would be upstream, and adult pallid sturgeon
would move farther than juvenile pallid sturgeon and shovelnose sturgeon.
Chapter three is a study on spawning migrations in shovelnose sturgeon. The
Missouri River hydrographs in 2008 and 2009 were partitioned into three hydrograph
phases based on the putative life-history and spawning requirements of shovelnose
sturgeon, and the environmental effect of water temperature was incorporated. The
specific objectives of chapter three were to (1) compare the movements of spawning
5
shovelnose sturgeon between years with differing spring hydrographs, (2) compare
movements among different reproductive categories, (3) document spawning locations of
female shovelnose sturgeon relative to larval drift distance, and (4) to document substrate
availability in the upper Missouri River and assess use relative to reproductive category.
I hypothesized that shovelnose sturgeon would make upstream movements to spawn,
movement of spawning fish would be greater in a year with higher discharge, and that
spawning fish would have greater movements than reproductively inactive fish.
Chapter four is a summary of major findings and a discussion integrating chapters
two and three. Implications for river management relative to Scaphirhynchus spp.
ecology are discussed.
6
References
Bailey, R. M., and F. B. Cross. 1954. River sturgeons of the American genus
Scaphirhynchus: characters, distribution, and synonymy. Papers of the Michigan
Academy of Science, Arts, and Letters 39:169-208.
Becker, G. C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison
Wisconsin.
Braaten, P. J., D. B Fuller, L. D. Holte, R. D. Lott, W. Viste, T. F. Brandt, and R. G.
Legare. 2008. Drift dynamics of larval pallid sturgeon and shovelnose sturgeon
in a natural side channel of the upper Missouri River, Montana. North American
Journal of Fisheries Management 28:808-826.
DeLonay, A. J., D. M. Papoulias, M. L. Wildhaber, M. L. Annis, J. L. Bryan, S. A.
Griffith, S. H. Holan, and D. E. Tillitt. 2007. Use of behavioral and
physiological indicators to evaluate Scaphirhynchus sturgeon spawning success.
Journal of Applied Ichthyology 23:428-435.
Dryer, M. P., and A. J. Sandvol. 1993. Pallid sturgeon recovery plan. U.S. Fish and
Wildlife Service, Bismarck, North Dakota.
Galat, D. L., C. R. Berry, W. M. Gardner, J. C. Hendrickson, G. E. Mestl, G. J. Power, C.
Stone, and M. R. Winston. 2005. Spatiotemporal patterns and changes in
Missouri River fishes. American Fisheries Society Symposium 45:249-291.
Galat, D. L., M. L. Wildhaber, and D. J. Dieterman. 2001. Spatial patterns of physical
habitat. Population structure and habitat use of benthic fishes along the Missouri
and lower Yellowstone Rivers, Vol. 2 US Geological Survey Cooperative
Research Units, University of Missouri, Columbia, Missouri. 91pp.
Gardner, W. M., and C. B. Jensen. 2011. Upper Missouri River basin pallid sturgeon
study 2005-2010 final report. United States Bureau of Reclamation, Montana
Department of Fish, Wildlife and Parks, Helena.
Gerrity, P. C., C. S. Guy, and W. M. Gardner. 2008. Habitat use of juvenile pallid
sturgeon and shovelnose sturgeon with implications for water level management
in a downstream reservoir. North American Journal of Fisheries Management
28:832-843.
Goodman, B. J. 2009. Ichthyoplankton density and shovelnose sturgeon spawning in
relation to varying discharge treatments. Master’s thesis. Montana State
University, Bozeman, Montana.
7
Hesse, L. W., J. C. Schmulbach, J. M. Carr, K. D. Keenlyne, D. G. Unkenholz, J. W.
Robinson, and G. E. Mestl. 1989. Missouri River fishery resources in relation to
past, present, and future stresses. Canadian Special Publication of Fisheries and
Aquatic Sciences 106:352-371.
Jager, H. I., W. Van Winkle, J. A. Challender, K. B. Lepla, P. Bates, and T. D. Counihan.
2002. A simulation study of factors controlling white sturgeon recruitment in the
Snake River. Pages 127-150 in W. Van Winkle, P. J. Andres, D. H. Secor, and D.
A. Dixon, editors. Biology, management and protection of North American
sturgeon. American Fisheries Society, Symposium 28, Bethesda, Maryland.
Junk, W. J., P. B. Bayley, and R. E. Sparks. 1989. The flood pulse concept in river
floodplain systems. Canadian Special Publication of Fisheries and Aquatic
Sciences 106:110-127.
Kallemeyn, L. W. 1983. Status of the pallid sturgeon (Scaphirhynchus albus). Fisheries
8:3-9.
Keenlyne, K. D. 1997. Life history and status of the shovelnose sturgeon,
Scaphirhynchus platorynchus. Environmental Biology of Fishes 48: 291-298.
Keenlyne, K. D., and L. G. Jenkins. 1993. Age at sexual maturity of the pallid sturgeon.
Transactions of the American Fisheries Society. 122:393-396.
Koch, J. D., and M. C. Quist. 2010. Current status and trends in shovelnose sturgeon
(Scaphirhynchus platorynchus) management and conservation. Journal of
Applied Ichthyology 26:491-498.
Poff, N. L., J. D. Allan, M. B. Bain, J. R. Karr, K. L. Prestegarrd, B. D. Richter, R. E.
Sparks, and J. C. Stromberg. 1997. The natural flow regime. Bioscience 47:769784.
Rogers, K. B., and G. C. White. 2007. Analysis of movement and habitat use from
telemetry data. Pages 625-676 in C. S. Guy, and M. L. Brown, editors. Analysis
and interpretation of freshwater fisheries data. American Fisheries Society,
Bethesda, Maryland.
Sparks, R. E., P. B. Bayley, S. L. Kohler, and L. L. Osborne. 1990. Disturbance and
recovery of large floodplain rivers. Environmental Management 14:699-709.
Stanford, J. A., J. V. Ward, W. J. Liss, C. A. Frissell, R. N. Williams, J. A. Lichatowich,
and C. C. Coutant. 1996. A general protocol for restoration of regulated rivers.
Regulated Rivers: Research and Management 12:391-413.
8
Tripp, S. J., Q. E. Phelps, R. E. Colombo, J. E. Garvey, B. M. Burr, D. P. Herzog, and R.
A. Hrabik. 2009. Maturation and reproduction of shovelnose sturgeon in the
middle Mississippi River. North American Journal of Fisheries Management
29:730-738.
USFWS (United States Fish and Wildlife Service). 2000. Biological opinion on the
operation of the Missouri River main stem reservoir system, operation and
maintenance of the Missouri River bank stabilization and navigation project,
and operation of the Kansas River reservoir system. USFWS, Bismarck, North
Dakota.
USFWS (United States Fish and Wildlife Service). 2003. Amendment to the 2000
biological opinion on the operation of the Missouri River main stem reservoir
system, operation and maintenance of the Missouri River bank stabilization and
navigation project, and operation of the Kansas River reservoir system. USFWS,
Minneapolis, Minnesota.
Webb, M. A. H., J. E. Williams, and L. R. Hildebrand. 2005. Recovery program review
for endangered pallid sturgeon in the upper Missouri River basin. Reviews in
Fisheries Science 13:165-176.
9
CHAPTER TWO
MOVEMENTS OF SCAPHIRHYNCHUS SPP. IN THE UPPER MISSOURI RIVER
Contributions of Authors and Co-Authors
Manuscript in Chapter 2
Author: Ryan R. Richards
Contributions: Conceived the study, collected and analyzed data, and authored the
manuscript.
Co-author: Christopher S. Guy
Contributions: Conceived the study, obtained funding and assisted with study design and
interpretation of analyses. Discussed the results and implications and edited the
manuscript.
Co-author: William M. Gardner
Contributions: Conceived the study, obtained funding, collected data, discussed the
results and implications.
10
Manuscript Information Page
Ryan R. Richards, Christopher S. Guy, William M. Gardner
Journal Name: Transactions of the American Fisheries Society
Status of Manuscript:
_x_Prepared for submission to a peer-reviewed journal
___Officially submitted to a peer-reviewed journal
___Accepted by a peer-reviewed journal
___Published in a peer-reviewed journal
Published by the American Fisheries Society
11
Abstract
Pallid sturgeon are endangered and shovelnose sturgeon populations, although
generally stable, are declining or extirpated in some areas. Altered discharge regimes and
habitat alteration and fragmentation caused by river regulation are among the primary
causes. Understanding how river regulation can affect movements of Scaphirhynchus
spp. is an important goal for sturgeon conservation. The objectives of this study were to
(1) compare the movements of juvenile pallid sturgeon, adult pallid sturgeon, and
shovelnose sturgeon among years with differing spring hydrographs, and (2) compare
movement patterns and distributions of juvenile pallid sturgeon, adult pallid sturgeon,
and shovelnose sturgeon locations in the upper Missouri River. Twenty-four hatcheryreared juvenile pallid sturgeon, seven adult pallid sturgeon, and 192 shovelnose sturgeon
were tracked from 2006 through 2009. No differences in movement rates were observed
for hatchery-reared juvenile pallid sturgeon, adult pallid sturgeon, or shovelnose sturgeon
among years despite four differing hydrographs representing a range of conditions from
below average to above average discharge with varying discharge rates of change, and
differing timing and duration of peak discharge. Adult pallid sturgeon had the most
extensive movements (194.9 km; 90% confidence interval, ± 69.6 km), followed by
shovelnose sturgeon (100.5 km; ± 12.6 km), and juvenile pallid sturgeon (38.2 km; ±
12.1 km). Some Scaphirhynchus spp. moved upstream, some downstream, and some did
not move. Shovelnose sturgeon were located an additional 126 km upstream of juvenile
pallid sturgeon locations, and an additional 94 km upstream from adult pallid sturgeon
12
locations, the majority of which were within 75 km of the headwaters of Fort Peck
Reservoir.
Introduction
Pallid sturgeon Scaphirhynchus albus are an endangered species (Dryer and
Sandvol 1993), and shovelnose sturgeon Scaphirhynchus platorynchus (hereafter referred
to collectively as Scaphirhynchus spp.) populations, while generally stable are declining
or extirpated in some areas (Koch and Quist 2010). Habitat alteration and fragmentation
due to dams has been implicated as the primary causes for declines in some
Scaphirhynchus spp. populations (Kallemeyn 1983; Dryer and Sandvol 1993; Keenlyne
1997). Shovelnose sturgeon have continued to successfully recruit in the Missouri River
downstream of Morony Dam near Great Falls, Montana, to the headwaters of Fort Peck
Reservoir (hereafter referred to as the upper Missouri River; Figure 2.1). Conversely, no
natural pallid sturgeon recruitment has been documented for at least 40 years, and
estimates of the number of wild adult pallid sturgeon in the upper Missouri River varied
from 40 to 100 in 2003 (Webb et al. 2005). However, artificial propagation and
reintroduction has resulted in juvenile pallid sturgeon that have been acclimated to the
conditions of the upper Missouri River for many years (Webb et al. 2005; Gerrity et al.
2008).
Most freshwater sturgeon species migrate within river systems, with large-bodied
species tending to move long distances (Auer 1996). Rising discharge associated with
the spring pulse is hypothesized to cue upstream migrations in Scaphirhynchus spp.
13
(Becker 1983; Keenlyne and Jenkins 1993; Goodman 2009; Tripp et al. 2009), and
regulation of the Missouri River has altered the environmental migration cue of spring
discharge (Galat et al. 2001). Pre-impoundment, the Missouri River hydrograph was
characterized by a large, highly variable spring discharge that is now more constant, and
reduced in both magnitude and duration (Hesse and Mestl 1993; Galat and Lipkin 2000;
Pegg et al. 2003). In the years following impoundment, discharge was reduced to
provide for flood control from April to July and increased from July to April to facilitate
hydropower production, irrigation, and downstream navigation (Pflieger and Grace
1987). However, based on the hypothesis that spring discharge is a spawning cue for
pallid sturgeon, the U.S. Fish and Wildlife Service (USFWS) has required that Fort Peck
and Gavins Point dams on the Missouri River be managed for increased spring discharge,
maintaining that previous spring discharges below these dams were not sufficient to
initiate spawning migrations in pallid sturgeon and other native fish (USFWS 2000,
2003).
Several dams exist in the Missouri River system upstream of the study area, with
three dams operated by the U.S. Bureau of Reclamation (USBR) having the most effect
on discharge and habitat (Gardner and Jensen 2011). These include Canyon Ferry Dam
on the Missouri River near Helena, Montana (rkm 3,626, measuring from the confluence
with the Mississippi River), Tiber Dam on the Marias River near Chester, MT (rkm 129,
measuring from confluence with the Missouri River), and Gibson Dam on the Sun River
(rkm 163, measuring from confluence with the Missouri River) (Gardner and Jensen
2011). Although the hydrograph is affected by the operation of these dams, due to
14
limited reservoir storage capacity, multi-use management of reservoirs, and numerous
unregulated tributaries that limit the influence of upstream impoundments, the upper
Missouri River is considered the least hydrologically altered section of the Missouri
River (Scott et al. 1997; Galat and Lipkin 2000; Pegg et al. 2003; Galat et al. 2005).
Operation of reservoirs may affect initiation of migrations by altering natural
discharge regimes, and affects the amount of habitat available for multiple
Scaphirhynchus spp. life history needs (Braaten et al. 2008; Gerrity et al. 2008).
Shovelnose sturgeon and pallid sturgeon larvae require long uninterrupted stretches of
free flowing riverine habitat for survival (Kynard et al. 2007; Braaten et al. 2008), and
dams on the Missouri River have fragmented habitat reducing the amount of unimpeded
river for drifting larvae (Braaten et al. 2008). White sturgeon Acipenser transmontanus
larvae require extended drift downstream of spawning areas and are likely to exhibit little
or no recruitment in reduced river reaches terminating in reservoirs (Jager et al. 2002).
Shovelnose sturgeon larvae may drift from 94 to 250 km, and pallid sturgeon larvae may
drift from 245 to 530 km after hatching, depending on water velocities (Braaten et al.
2008). The extended larval drift observed in pallid sturgeon may be contributing to the
lack of recruitment; conversely, larval shovelnose sturgeon have a shorter drift, and may
settle out of the water column before encountering reservoir headwaters, explaining the
disparity in recruitment between the two species.
Populations of wild pallid sturgeon in the upper Missouri River are nearly
extirpated (Webb et al. 2005). As hatchery-reared juveniles that have been released in
the wild approach reproductive age, understanding how river regulation affects
15
movements of pallid sturgeon becomes an important goal for conservation. The purpose
of this study was to document and compare movements of Scaphirhynchus spp. through
the annual spring pulse of the upper Missouri River, and assess use of the available
riverine habitat relative to larval drift distance. The specific objectives were to (1)
compare the movements of juvenile pallid sturgeon, adult pallid sturgeon, and shovelnose
sturgeon among years with differing spring hydrographs, and (2) compare movement
patterns and distributions of juvenile pallid sturgeon, adult pallid sturgeon, and
shovelnose sturgeon locations in the upper Missouri River. Based on what is currently
known and hypothesized of Scaphirhynchus spp. ecology, I hypothesized that movements
would be greater in a year with higher discharge, the highest movement rates would be
coincident to rising discharge, movements would be upstream, and due to larger body
size, adult pallid sturgeon would move farther than juvenile pallid sturgeon and
shovelnose sturgeon.
Methods
Fish Capture and Surgery
From 2006 to 2009 hatchery-reared juvenile pallid sturgeon, as described by
Gerrity et al. (2008), wild adult pallid sturgeon, and wild shovelnose sturgeon (hereafter
referred to as juvenile pallid sturgeon, adult pallid sturgeon, and shovelnose sturgeon,
respectively) were captured and implanted with radio transmitters in the upper Missouri
River. The Scaphirhynchus spp. used in this study were part of a larger study
investigating seasonal fish migrations in the upper Missouri River; thus, capture locations
16
and techniques varied and are detailed in Gardner and Jensen (2011). Juvenile pallid
sturgeon were selected for transmitter implantation if they were ≥ 0.68 kg, all adult pallid
sturgeon captured were implanted with transmitters, and shovelnose sturgeon were
implanted with transmitters if they were ≥ 1.4 kg. Transmitters implanted in
Scaphirhynchus spp. were selected to be as light as possible while maximizing battery
life and transmitter weight did not exceed 2% of fish body weight (Winter 1996).
Transmitter-implantation surgery was performed using methods modified from Ross and
Kleiner (1982) and Summerfelt and Smith (1990). All fish were placed in a holding tank
for 20 minutes to allow for recovery from the surgery, and were released near the capture
location. Sample sizes of juvenile pallid sturgeon, adult pallid sturgeon, and shovelnose
sturgeon for each year are found in Table 2.1.
Tracking
Individual fish were tracked weekly from May through August 2006-2009 to
estimate the direction and extent of movements associated with the spring pulse of the
hydrograph. Telemetered Scaphirhynchus spp. were located using a portable SRX-400
Lotek receiver (Lotek Wireless, Inc., Newmarket, Ontario, Canada) equipped with a boatmounted, four-element Yagi antenna. Precise locations were determined by triangulation
or by passing the boat directly over the fish using a three-element hand held Yagi antenna
(Winter 1996; Blanchfield et al. 2005); this method has been shown to be accurate to
within five meters by means of blind tests using transmitters placed in the river (Gerrity
et al. 2008; Bellgraph et al. 2008). Latitude, longitude, and river kilometer (rkm) were
measured at each Scaphirhynchus spp. location using a boat mounted GPS unit. Data
17
from eight autonomous land-based radio-receiving stations (Figure 2.1) were also
incorporated into the data set. These stations were located on the bank adjacent to the
river and consisted of a continuously recording solar powered SRX-400 Lotek receiver
attached to two four-element Yagi antennas.
Hydrographs
Hydrographs were derived by averaging the mean daily discharge recorded at
three USGS gauging stations on the Missouri River within the study site: Fort Benton
(rkm 3,337), Virgelle Ferry (rkm 3,274), and Landusky (rkm 3,090). For comparison to
average discharge, data from the same three USGS gauging stations were used to
calculate the mean daily discharge averaged over the years 1935 to 2005. Because rising
spring discharge is believed to initiate migrations in Scaphirhynchus spp. the hydrograph
for each year was partitioned into three phases. This allowed for comparison of
Scaphirhynchus spp. movements between periods of increasing discharge, decreasing
discharge, and baseflow. Hydrograph phases include ascending limb, descending limb,
and post-descending limb. The ascending limb for each year was defined as the period
from 1 May (arbitrarily selected based on when sampling started) to the peak of the
hydrograph, the descending limb was defined as the period post-peak until baseflow was
achieved, and the post-descending limb was defined as the period after baseflow was
achieved until tracking ceased on 31 August (Figure 2.2). Baseflows for each
hydrograph were derived using the constant-discharge baseflow separation method
(McCuen 2005).
18
Movement and Movement Rates
Mean daily total and net movement rates were calculated for each fish for each
phase of the hydrograph. Total daily movement rate was calculated for a given fish by
dividing the distance in river kilometers between successive relocations by the number of
days elapsed between relocations (White and Garrott 1990). Net daily movement rate
was calculated by dividing the difference in river kilometer between successive
relocations by the number of days that elapsed between relocations such that a positive
rate indicated upstream movement, a negative rate indicated downstream movement, and
a rate of zero indicated no movement (Bramblett 1996). Additional movement likely
occurs between relocations; thus, all movement rate data represent a minimum for the
period between relocations (Rogers and White 2007).
Repeated measures (with individual fish as the repeated variable) analysis of
variance (ANOVA) was used to test the hypotheses that there were no differences in net
and total movement rates of juvenile pallid sturgeon, adult pallid sturgeon, and
shovelnose sturgeon among 2006-2009. If movement rates did not differ significantly,
years were pooled to increase statistical power and repeated measures ANOVA was used
to test the hypotheses that there were no differences in total and net movement rates
among hydrograph phases. Bonferroni multiple-comparisons test was used to control the
experimentwise error rate (Rogers and White 2007). Individual radio-tagged fish were
the experimental unit for all statistical tests, and normality for all variables was assessed
using residual and normal probability plots. To reduce the chance of making a type II
error, an α = 0.1 was established a priori for all repeated measures ANOVAs. Sample
19
sizes of juvenile pallid sturgeon and adult pallid sturgeon were relatively small; however,
the repeated measures ANOVA used to analyze the data takes into consideration the
variation within individuals, thus offering high statistical power relative to sample size
(Ott 1993; Kutner et al. 2004).
Because fish were widely distributed throughout the study area when tracking
commenced in the spring, plots of river kilometer-standardized movement patterns were
made for each fish to graphically compare movements between pallid sturgeon and
shovelnose sturgeon among years. River kilometer at the first annual location of each
fish was standardized to zero with subsequent locations indicating the distance upstream
or downstream from the first location.
Results
Hydrographs
The ascending limb of the 2008 Missouri River hydrograph had a peak discharge
that was 45-116 % higher and a mean daily rate of change 3.3 to 5.2 times higher than the
remaining years (Figure 2.2). The ascending limb of the 2007 hydrograph had the lowest
peak discharge and lowest mean daily rate of change of all four years. Discharge during
the descending limb was greatest in 2008; however, mean daily rate of change was
similar for all four years. Post-descending limb discharge and mean daily rate of change
was similar for all four years (Table 2.2; Figure 2.2). In addition to higher peak
discharge achieved at a higher rate, discharge levels remain high for a greater duration in
2008 than in the remaining years. For example, after 2008 the highest peak discharge,
20
578 m3/s, was observed in 2009; in 2008 discharge remained above 578 m3/s for 28 days,
while in the remaining years discharge began to decrease sharply immediately after the
peaks were achieved (Figure 2.2).
The Missouri River hydrographs in 2006-2009 represent a range of conditions
relative to average discharge, including above and below average discharge years. When
compared to the average, discharge in 2008 was an above average discharge year, while
peak discharge in 2007 was below the 1935-2005 average; peak discharge in 2006 and
2009 were between these two extremes (Figure 2.2). The timing of peak discharge in
2006 and 2007 coincide with the peak in average discharge, while peak discharge in 2008
and 2009 was 2-3 weeks earlier (Figure 2.2). Both the timing and magnitude of peak
discharge in 2006 closely approximated that of the average; however in 2006 the duration
of the peak was much shorter than the average, a trend that is seen for all years except
2008. In the years 2006, 2007, and 2009 discharge remained elevated for a shorter
duration than the average, with discharge dropping to pre-peak levels quickly compared
to the more prolonged drop in discharge seen in the average. In 2008, discharge
remained elevated for a prolonged period, resembling the duration of increased discharge
seen in the average (Figure 2.2).
Movement and Movement Rates
Mean net (F3, 20 = 1.12, P = 0.36) and mean total (F3, 20 = 1.76, P = 0.19)
movement rates of juvenile pallid sturgeon did not differ significantly among years
within hydrograph phases (Table 2.3). When years were pooled, mean net (F2, 24 = 0.84,
P = 0.44) and total (F2, 24 = 0.58, P = 0.57) movement rates of juvenile pallid sturgeon did
21
not differ significantly among hydrograph phases (Table 2.3). Mean net (F3, 3 = 1.30, P =
0.42) and mean total (F3, 3 = 1.05, P = 0.49) movement rates of adult pallid sturgeon did
not differ significantly among years within hydrograph phases (Table 2.4). When years
were pooled, mean net (F2, 7 = 3.30, P = 0.10) and total (F2, 7 = 3.36, P = 0.10) movement
rates of adult pallid sturgeon differed significantly among hydrograph phases; however,
when the Bonferroni multiple-comparisons test was used to control the experimentwise
error rate there were no significant differences in net or total movement rates of adult
pallid sturgeon among hydrograph phases (P > 0.10) (Table 2.4). Mean net (F3, 188 =
1.00, P = 0.40) and mean total (F3, 188 = 1.46, P = 0.23) movement rates of shovelnose
sturgeon did not differ significantly among years within hydrograph phases (Table 2.5).
When years were pooled, mean net movement rate (F2, 199 = 0.57, P = 0.57) did not differ
significantly, but total movement rate (F2, 199 = 2.41, P = 0.09) differed significantly
among hydrograph phases. However, when the Bonferroni multiple-comparisons test
was used to control the experimentwise error rate there were no significant differences in
total movement rates of shovelnose sturgeon among hydrograph phases (P > 0.10) (Table
2.5). Scaphirhynchus spp. movements did not differ despite a range of annual discharge
from above to below the average, and movements were the same whether the hydrograph
was ascending, descending, or at baseflow.
The average sum of annual movements was greater for adult pallid sturgeon
(194.9 km; 90% confidence interval, ± 69.6 km), than juvenile pallid sturgeon (38.2 km;
± 12.1 km), and shovelnose sturgeon (100.5 km; ± 12.6 km). However, sample size of
adult pallid sturgeon (N = 7) was small compared to shovelnose sturgeon (N = 192), and
22
adult pallid sturgeon movements lacked low values, with a minimum of 77.5 km and a
maximum of 392.0 km. Conversely, shovelnose sturgeon movements were more variable
with a minimum of 0.5 km and a maximum of 516.4 km. Juvenile pallid sturgeon did not
exhibit extensive upstream or downstream movements through the four years examined,
while adult pallid sturgeon were observed moving upstream or downstream (Figure 2.3).
Shovelnose sturgeon movements were highly variable with the majority of fish remaining
sedentary and others moving extensively upstream or downstream (Figure 2.4).
Shovelnose sturgeon used more of the available riverine habitat than juvenile pallid
sturgeon or adult pallid sturgeon, with shovelnose sturgeon locations distributed across
298 km of the study area, while juvenile pallid sturgeon and adult pallid sturgeon
locations were distributed across 177 km and 191 km of the study area, respectively
(Figure 2.5). The highest densities of adult pallid sturgeon and juvenile pallid sturgeon
locations were in the 75 km upstream of the Fort Peck Reservoir headwaters, while
shovelnose sturgeon locations were distributed more evenly throughout the study area.
Shovelnose sturgeon locations were distributed 126 km farther upstream than juvenile
pallid sturgeon, and 94 km farther upstream than adult pallid sturgeon; the distribution of
downstream locations was similar for all three groups (Figure 2.5).
Discussion
I rejected my hypothesis that Scaphirhynchus spp. movements would be greater in
a year with higher discharge. Scaphirhynchus spp. movement rates did not differ among
years 2006-2009, despite four differing hydrographs from below average to above
23
average discharge, varying discharge rates of change, and differing timing and duration
of peak discharge. Similar results were observed in shovelnose sturgeon in the Kansas
River wherein movements were not related to discharge (Quist et al. 1999). Rising
discharge has been proposed as a cue initiating spring migrations in Scaphirhynchus spp.,
and movement is believed to be upstream (Becker 1983; Keenlyne and Jenkins 1993;
Goodman 2009; Tripp et al. 2009). However, the hypotheses that movement rates would
be greatest when discharge was rising and that movements would be upstream were
rejected. No differences were found in Scaphirhynchus spp. movement rates among
periods of rising discharge, decreasing discharge, and baseflow, and Scaphirhynchus spp.
in the upper Missouri River were observed not moving, moving upstream, and moving
downstream through the course of the spring hydrograph. I failed to reject the hypothesis
that adult pallid sturgeon would move farther than juvenile pallid sturgeon and
shovelnose sturgeon. On average adult pallid sturgeon moved five times as far as
juvenile pallid sturgeon, and nearly twice as far as shovelnose sturgeon.
Most spring migrations in sturgeon are believed to be upstream (Hall et al. 1991;
Bemis and Kynard 1997; Wilson and McKinley 2004). However, no movement,
movements upstream, and movements downstream were observed in adult pallid
sturgeon and shovelnose sturgeon in the upper Missouri River. Adult pallid sturgeon also
moved upstream and downstream during the spring pulse in the Missouri River below
Fort Randall Dam and in Lake Sharpe (Erickson 1992; Wanner et al. 2007), and 60% of
shovelnose sturgeon in the Yellowstone and Missouri rivers below Fort Peck Reservoir
had winter locations upstream of summer locations (Bramblett and White 2001).
24
Scaphirhynchus spp. in the upper Missouri River are isolated from downstream
populations and bounded by Fort Peck Reservoir; however the current upstream
distributions of pallid sturgeon and shovelnose sturgeon in the upper Missouri River
closely match historical distributions (Wilson and McKinley 2004). Scaphirhynchus spp.
in the upper Missouri River cannot make extensive upstream movements without
encountering unsuitable water temperatures and reduced turbidity gradients, and
Scaphirhynchus spp. attempting to move downstream encounter unsuitable habitat in the
form of the headwaters of Fort Peck Reservoir.
On average, adult pallid sturgeon moved longer distances than juvenile pallid
sturgeon or shovelnose sturgeon. This was expected as pallid sturgeon had higher
movement rates than shovelnose sturgeon in the Yellowstone and Missouri rivers below
Fort Peck Reservoir (Bramblett and White 2001), and the distances covered during spring
migrations by sturgeon often have a positive relationship with average adult body size
(Auer 1996). Although adult pallid sturgeon moved farther on average than shovelnose
sturgeon, shovelnose sturgeon locations were distributed farther upstream than adult
pallid sturgeon locations. Pallid sturgeon larvae may drift for 11-17 dph and cover from
245 to 530 km; shovelnose sturgeon larvae may drift for 6 days post hatch (dph) and
cover distances from 94 to 250 km (Braaten et al. 2008). The extended duration and
distance of pallid sturgeon larval drift is significant when considering that there is only
about 380 km of riverine habitat between Morony Dam and the headwaters of Fort Peck
Reservoir. Although sex and reproductive stage were unknown, the farthest upstream
adult pallid sturgeon location in the four years of this study was approximately 250 km
25
upstream of the Fort Peck Reservoir headwaters, with the majority of locations in the 75
km upstream of the headwaters. Conversely, the farthest upstream shovelnose sturgeon
locations were approximately 350 km above the headwaters. The extended larval drift
observed in pallid sturgeon may be contributing to the lack of recruitment observed in the
upper Missouri River (Braaten et al. 2008), and is cause for increased study.
Scaphirhynchus spp. are lotic and require riverine habitat through all life stages
(Bailey and Cross 1954; Kallemeyn 1983; Keenlyne 1997; Wildhaber et al. 2011).
Previous telemetry studies have shown that Scaphirhynchus spp. in the upper Mississippi
River, the upper Missouri River, and the Yellowstone and Missouri rivers above Lake
Sakakawea generally avoided impounded areas (Curtis et al. 1997; Bramblett and White
2001; Gerrity et al. 2008). Manipulation of the operational pool level of Fort Peck
Reservoir can influence the amount of available Scaphirhynchus spp. habitat through
inundation of lotic habitat as the operational pool level rises, and Scaphirhynchus spp.
have been shown to range further downstream in the upper Missouri River as the
operational pool level of Fort Peck Reservoir declines (Gerrity et al. 2008). Fort Peck
Reservoir operational pool levels in 2004 made available an additional 60 km of lotic
habitat that would have been inundated if the reservoir was at full operational pool level
(Gerrity et al. 2008); thus, implications exist for river management as the highest
densities of juvenile pallid sturgeon and adult pallid sturgeon locations were in the 75 km
directly upstream from the Fort Peck Reservoir headwaters.
Movements of reproductively inactive female shovelnose sturgeon were
significantly less than those of male, and potential spawners, and confirmed spawning
26
females in the upper Missouri River (Chapter 3), significantly reduced movement in
reproductively inactive adult shovelnose sturgeon compared to spawners was observed in
the lower Missouri River (DeLonay et al. 2009), and juvenile pallid sturgeon in this study
moved less than adult pallid sturgeon. It is likely that disregarding the effects of
maturity, sex, and reproductive stage on movements of Scaphirhynchus spp. while
conducting telemetry studies is resulting in highly variable movement data and indicates
the need for identification of sex and reproductive stage of study subjects.
Increasingly, studies are failing to find a link between discharge and initiation of
migration in Scaphirhynchus spp. (DeLonay et al. 2009; Papoulias et al. 2011). Rather
than discharge, it is believed that reproductive maturation in Scaphirhynchus spp. is
initiated many months prior to spawning, with day length as the environmental cue, while
spawning is initiated by the proximate cue of water temperature (DeLonay et al. 2009;
Papoulias et al. 2011; Wildhaber et al. 2011). Movement rates of confirmed spawning
female shovelnose sturgeon in the upper Missouri River peaked at water temperatures
from 18 to 22 °C regardless of discharge, indicating that water temperature is a more
likely proximate cue initiating spawning migrations (Chapter 3). In this study, no
differences were observed in movement rates despite four hydrographs representing
conditions from above to below average discharge, with variable rates of change and
different timing and duration of peak discharge; thus, providing further evidence that
discharge does not influence movements in Scaphirhynchus spp. Although the variations
in discharge examined in this study did not affect movements of Scaphirhynchus spp.,
discharge may be important in determining spawning success and recruitment. Prolonged
27
high discharge has been shown to cause protracted spawning and a greater abundance of
larval and age-0 Scaphirhynchus spp. (Goodman 2009; Phelps et al. 2010); how
discharge influences recruitment in the upper Missouri River is unknown and is cause for
further study.
Acknowledgements
The U.S. Bureau of Reclamation and PPL Montana provided funding and support
for this research. I thank Robert Bramblett for manuscript reviews; Eli Cureton, Joel Van
Eenennaam, Casey Jensen, Eli McCord, Mike Wente, Randy Rodencal, Ben Cox, Matt
Jaeger, Michael Michelson, Nick Smith, Sue Camp, Steve Leathe, Mike Meeuwig, Ben
Goodman, Boone Richards, Justin Spinelli, John Syslo, Peter Brown, Steven Ranney, and
Mariah Talbott for their invaluable assistance in the field, lab, and office. The Montana
Cooperative Fisheries Research Unit is jointly sponsored by the Montana Fish, Wildlife,
and Parks, Montana State University, and the U. S. Geological Survey.
28
Tables
Table 2.1. Sample sizes for juvenile pallid sturgeon, adult pallid sturgeon, and
shovelnose sturgeon in the Missouri River above Fort Peck Reservoir 2006-2009.
Year
2006
2007
2008
2009
Juvenile pallid
sturgeon
7
8
5
4
Species
Adult pallid
sturgeon
2
1
2
2
Shovelnose
sturgeon
28
51
55
58
29
Table 2.2. Minimum, maximum, and mean (SD) daily discharge (m3/s) and daily
discharge rate of change (ROC) (m3/s/d) for the Missouri River upstream of Fort Peck
Reservoir for all hydrograph phases, 2006- 2009. See text for phase definitions.
Variable
Year
Phase
Daily discharge
2006
2007
2008
2009
Daily discharge ROC
2006
2007
2008
2009
Minimum
Maximum
Mean (SD)
Ascending
Descending
Post-descending
Ascending
Descending
Post-descending
Ascending
238
152
141
205
152
120
145
546
506
163
389
373
149
839
292 (59)
230 (106)
151 (7)
271 (52)
208 (58)
131 (8)
259 (192)
Descending
Post-descending
196
156
823
196
515 (213)
183 (10)
Ascending
Descending
299
194
578
573
436 (76)
313 (77)
Post-descending
161
216
184 (14)
Ascending
Descending
Post-descending
Ascending
Descending
Post-descending
-32
-39
-8
-19
-39
-9
120
5
8
49
3
6
6 (28)
-10 (13)
0 (3)
5 (15)
-7 (8)
-1 (3)
Ascending
Descending
Post-descending
Ascending
Descending
Post-descending
-5
-65
-11
-14
-48
-18
165
45
6.00
39
36
18
26 (46)
-9 (20)
-1 (4)
8 (17)
-7 (17)
-1 (8)
30
Table 2.3. Minimum, maximum, and mean ± 90% confidence interval (CI) net and total
movement rates (km/d) for juvenile pallid sturgeon in the Missouri River upstream of
Fort Peck Reservoir for all hydrograph phases 2006-2009. Positive net movement rates
represent overall upstream movement, negative rates represent downstream movement.
See text for hydrograph phase definitions.
Variable
Net movement rate
Year
2006
2007
2008
2009
Pooled
Total movement rate
2006
2007
2008
2009
Pooled
Phase
Minimum
Maximum
Mean ± 90% CI
Ascending
-0.92
0.04
-0.21 ± 0.21
Descending
-0.81
0.26
-0.08 ± 0.22
Post-descending
-0.06
0.08
0.00 ± 0.03
Ascending
-0.08
0.34
0.02 ± 0.08
Descending
-1.25
0.27
-0.13 ± 0.27
Post-descending
-0.16
0.18
-0.01 ± 0.06
Ascending
-0.88
0.89
0.04 ± 0.46
Descending
-0.57
0.29
-0.15 ± 0.26
Post-descending
-0.24
0.29
0.03 ± 0.18
Ascending
-0.04
0.21
0.04 ± 0.10
Descending
-0.08
0.42
0.09 ± 0.18
Post-descending
-0.98
0.02
-0.44 ± 0.48
Ascending
-0.92
0.89
-0.04 ± 0.12
Descending
-1.25
0.42
-0.08 ± 0.12
Post-descending
-0.98
0.29
-0.06 ± 0.09
Ascending
0.09
1.34
0.55 ± 0.27
Descending
0.04
0.81
0.28 ± 0.16
Post-descending
0.02
0.13
0.07 ± 0.02
Ascending
0.03
0.34
0.09 ± 0.06
Descending
0.01
1.65
0.32 ± 0.32
Post-descending
0.00
0.92
0.24 ± 0.17
Ascending
0.04
0.89
0.45 ± 0.31
Descending
0.11
0.86
0.47 ± 0.22
Post-descending
0.00
0.29
0.15 ± 0.11
Ascending
0.00
0.21
0.07 ± 0.07
Descending
0.04
0.88
0.38 ± 0.31
Post-descending
0.23
0.98
0.53 ± 0.38
Ascending
0.00
1.34
0.30 ± 0.12
Descending
0.01
1.65
0.35 ± 0.13
Post-descending
0.00
0.98
0.21 ± 0.09
31
Table 2.4. Minimum, maximum, and mean ± 90% confidence interval (CI) net and total
movement rates (km/d) for adult pallid sturgeon in the Missouri River upstream of Fort
Peck Reservoir for all hydrograph phases 2006-2009. Positive net movement rates
represent overall upstream movement, negative rates represent downstream movement.
See text for hydrograph phase definitions.
Variable
Net movement rate
Year
2006
2007
2008
2009
Pooled
Total movement rate
2006
2007
2008
2009
Pooled
Phase
Minimum
Maximum
Mean ± 90% CI
Ascending
2.38
8.39
5.38 ± 4.95
Descending
-1.69
-0.34
-1.02 ± 1.11
Post-descending
0.28
0.78
0.53 ± 0.41
Ascending
-0.69
-0.69
-0.69
Descending
-2.60
-2.60
-2.60
Post-descending
-0.04
-0.04
-0.04
Ascending
-0.17
-0.10
-0.13 ± 0.05
Descending
-1.48
-0.29
-0.89 ± 0.98
Post-descending
0.00
0.00
0.00
Ascending
-2.04
1.89
-0.07 ± 3.23
Descending
-2.29
0.40
-0.94 ± 2.21
Post-descending
-0.57
0.01
-0.28 ± 0.48
Ascending
-2.04
8.39
1.38 ± 2.14
Descending
-2.60
0.40
-1.18 ± 0.70
Post-descending
-0.57
0.78
0.08 ± 0.30
Ascending
2.38
8.47
5.42 ± 5.01
Descending
1.02
4.09
2.55 ± 2.52
Post-descending
0.28
0.78
0.53 ± 0.41
Ascending
1.28
1.28
1.28
Descending
2.60
2.60
2.60
Post-descending
0.68
0.68
0.68
Ascending
0.39
0.50
0.45 ± 0.09
Descending
1.64
2.03
1.83 ± 0.32
Post-descending
0.00
0.00
0.00
Ascending
1.89
4.12
3.01 ± 1.84
Descending
2.78
2.79
2.79 ± 0.01
Post-descending
0.01
2.90
1.45 ± 1.45
Ascending
0.39
8.47
2.72 ± 1.76
Descending
1.02
4.09
2.42 ± 0.61
Post-descending
0.00
2.90
0.78 ± 0.73
32
Table 2.5. Minimum, maximum, and mean ± 90% confidence interval (CI) net and total
movement rates (km/d) for shovelnose sturgeon in the Missouri River upstream of Fort
Peck Reservoir for all hydrograph phases 2006-2009. Positive net movement rates
represent overall upstream movement, negative rates represent downstream movement.
See text for hydrograph phase definitions.
Variable
Net movement rate
Year
2006
2007
2008
2009
Pooled
Total movement rate
2006
2007
2008
2009
Pooled
Phase
Minimum
Maximum
Mean ± 90% CI
Ascending
-1.16
3.96
0.04 ± 0.31
Descending
-4.65
3.40
0.21 ± 0.53
Post-descending
-5.37
2.59
0.22 ± 0.44
Ascending
-3.37
3.27
-0.10 ± 0.25
Descending
-2.44
4.01
0.27 ± 0.22
Post-descending
-4.63
3.03
0.07 ± 0.22
Ascending
-5.06
4.51
0.15 ± 0.30
Descending
-1.71
2.31
0.13 ± 0.17
Post-descending
-2.45
1.94
0.02 ± 0.15
Ascending
-5.44
4.46
-0.27 ± 0.35
Descending
-3.51
2.96
-0.02 ± 0.17
Post-descending
-5.68
8.29
0.04 ± 0.44
Ascending
-5.44
4.51
-0.05 ± 0.16
Descending
-4.65
4.01
0.13 ± 0.12
Post-descending
-5.68
8.29
0.07 ± 0.16
Ascending
0.03
4.40
1.07 ± 0.33
Descending
0.02
7.16
1.23 ± 0.53
Post-descending
0.00
5.37
0.81 ± 0.38
Ascending
0.00
11.93
1.22 ± 0.45
Descending
0.01
4.81
0.87 ± 0.25
Post-descending
0.00
4.91
0.59 ± 0.21
Ascending
0.01
5.06
0.78 ± 0.26
Descending
0.01
3.40
0.82 ± 0.18
Post-descending
0.00
2.45
0.39 ± 0.12
Ascending
0.01
5.49
1.06 ± 0.32
Descending
0.00
6.18
1.09 ± 0.28
Post-descending
0.00
8.29
1.06 ± 0.39
Ascending
0.00
11.93
1.02 ± 0.18
Descending
0.00
7.16
0.97 ± 0.14
Post-descending
0.00
8.29
0.71 ± 0.15
33
Figures
Figure 2.1. Map of Missouri and Marias rivers in Montana, with study area in detail.
34
900
AL
DL
PDL
AL
DL
PDL
750
600
450
Mean daily discharge (m3/s)
300
150
2006
0
May
Jun
Jul
Aug
Sep
2007
May
Jun
Jul
Aug
Sep
Discharge
Average discharge
900
AL
DL
PDL
AL
DL
PDL
750
600
450
300
150
2008
0
May
Jun
Jul
Month
Aug
Sep
2009
May
Jun
Jul
Aug
Sep
Month
Figure 2.2. Hydrograph phases for the Missouri River upstream of Fort Peck Reservoir,
Montana 2006-2009, as defined by mean daily discharge. Average discharge (dashed
line) represents the mean daily discharge averaged over 1935-2005. Phases include
ascending limb (AL), descending limb (DL), and post-descending limb (PDL). See text
for phase definitions.
35
250
AL
DL
PDL
AL
DL
PDL
125
Standardized distance (km)
0
-125
2006
-250
May
Jun
Jul
Aug
Sep
2007
May
Jun
Jul
Aug
Sep
Juvenile
Adult
250
AL
DL
PDL
AL
DL
PDL
125
0
-125
2008
-250
May
Jun
Jul
Month
Aug
Sep
2009
May
Jun
Jul
Aug
Sep
Month
Col 1 vs Col 2
Col 4 vs Col 5
Col 16 vs Col 8
Col 10 vs Col 11
Col 13 vs Col 14
Col 16 vs Col 17
Col 19 vs Col 20
Col 22 vs Col 23
Figure 2.3. Plots of river kilometer-standardized movement patterns of juvenile pallid
sturgeon (solid lines) and adult pallid sturgeon (dashed lines) in the Missouri River above
Fort Peck Reservoir, Montana 2006-2009. The river kilometer of the first location of
each fish has been standardized to zero; values above zero represent upstream movement,
and values below zero represent downstream movement. Hydrograph phases include
ascending limb (AL), descending limb (DL), and post-descending limb (PDL). See text
for phase definitions.
36
250
AL
PDL
DL
AL
DL
PDL
125
Standardized distance (km)
0
-125
2006
-250
May
250
2007
Jun
AL
Jul
Aug
Sep
May
PDL
DL
Jun
AL
Jul
Aug
Sep
PDL
DL
125
0
-125
2008
-250
May
2009
Jun
Jul
Month
Aug
Sep
May
Jun
Jul
Aug
Sep
Month
Figure 2.4. Plots of river kilometer-standardized movement patterns of shovelnose
sturgeon in the Missouri River above Fort Peck Reservoir, Montana 2006-2009. The
river kilometer of the first location of each fish has been standardized to zero; values
above zero represent upstream movement, and values below zero represent downstream
movement. Hydrograph phases include ascending limb (AL), descending limb (DL), and
post-descending limb (PDL). See text for phase definitions.
37
Figure 2.5. Distributions of juvenile pallid sturgeon, adult pallid sturgeon, and
shovelnose sturgeon in the Missouri River above Fort Peck Reservoir, Montana 20062009.
38
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43
CHAPTER THREE
SPAWNING RELATED MIGRATION OF SHOVELNOSE STURGEON IN THE
MISSOURI RIVER ABOVE FORT PECK RESERVOIR, MONTANA
Contributions of Authors and Co-Authors
Manuscript in Chapter 3
Author: Ryan R. Richards
Contributions: Conceived the study, collected and analyzed data, and authored the
manuscript.
Co-author: Christopher S. Guy
Contributions: Conceived the study, obtained funding, and assisted with study design and
interpretation of analyses. Discussed the results and implications and edited the
manuscript.
Co-author: Molly A. Webb
Contributions: Conceived the study, provided support and training for histological and
physiological analyses. Assisted with interpretation of analyses, discussed the results and
implications.
Co-author: William M. Gardner
Contributions: Conceived the study, obtained funding, collected data, discussed the
results and implications.
Co-author: Casey B. Jensen
Contributions: Collected data, assisted with analyses, and discussed the results and
implications.
44
Manuscript Information Page
Ryan R. Richards, Christopher S. Guy, Molly A. Webb, William M. Gardner, Casey B.
Jensen
Journal Name: Transactions of the American Fisheries Society
Status of Manuscript:
_x_Prepared for submission to a peer-reviewed journal
___Officially submitted to a peer-reviewed journal
___Accepted by a peer-reviewed journal
___Published in a peer-reviewed journal
Published by the American Fisheries Society
45
Abstract
Some shovelnose sturgeon populations are extirpated or in decline, with reservoir
operation implicated as a primary cause. Understanding how river regulation affects
movements of spawning shovelnose sturgeon is imperative for conservation. The
hypotheses of this study were (1) that shovelnose sturgeon would make upstream
movements to spawn, (2) movement of spawning fish would be greater in a year with
higher discharge, and (3) that spawning fish would have greater movements than
reproductively inactive fish. Seventy-eight shovelnose sturgeon in five reproductive
categories (e.g., males, confirmed spawning females, potentially spawning females,
atretic females, and reproductively inactive females) were tracked in 2008 and 2009. No
differences in movement rates were observed in confirmed spawning females between
years although in 2008 peak flows were obtained at a faster rate, flows were higher by
45%, and high discharge was of a greater duration. Reproductively inactive females were
the only category to differ in movements and did not have extensive movements. All
other reproductive categories, including confirmed spawning females, exhibited extensive
migrations. Movements of fish in all reproductive categories were variable, with some
fish migrating upstream, some downstream, and some not migrating. The highest
movement rates of confirmed spawning females were at temperatures between 18 and 22
°C regardless of discharge, indicating that water temperature is a more likely proximate
cue initiating spawning migrations in shovelnose sturgeon.
46
Introduction
Many aquatic species have evolved life-history strategies in direct response to
natural flow regimes, with flow linked to reproduction, larval survival, growth, and
recruitment (Bunn and Arthington 2002). Shovelnose sturgeon Scaphirhynchus
platorynchus have been indigenous to the large, turbid rivers of the Missouri and
Mississippi River basins for millions of years (Bailey and Cross 1954) and have a lifehistory strategy that is dependent on the conditions of the environment (Wildhaber et al.
2011). Dams and impoundments can fragment fish habitat, as well as alter and decouple
natural discharge and water temperature regimes, reduce sediment transport, and
disconnect the floodplain from the river channel (Junk et al. 1989; Sparks et al. 1990;
Stanford et al. 1996; Poff et al. 1997). Although most shovelnose sturgeon populations
are stable, some are in decline or extirpated (Koch and Quist 2010). Habitat alteration
and fragmentation due to dams are implicated as primary causes (Keenlyne 1997).
Changes in discharge and water temperature associated with the spring pulse are
believed to cue spawning migrations in adult sturgeons (Becker 1983; Keenlyne and
Jenkins 1993; DeLonay et al. 2009). Reproductively active sturgeon can fail to spawn if
the environmental cues of discharge and water temperature are absent (Auer 2004), and
in the Marias River, a tributary to the main stem Missouri River, shovelnose sturgeon
have been shown to initiate spawning only when a threshold discharge was reached
(Goodman 2009). The construction and operation of dams on the Missouri River has
altered and decoupled the putative environmental spawning cues of discharge and water
temperature thought to initiate spawning in shovelnose sturgeon (Galat et al. 2001). Prior
47
to dam construction, the Missouri River hydrograph was characterized by a large, highly
variable spring pulse that is now more constant and reduced in magnitude and duration
(Hesse and Mestl 1993; Galat and Lipkin 2000; Pegg et al. 2003).
Shovelnose sturgeon larvae drift for six days post-hatch, and require long
uninterrupted stretches of free flowing riverine habitat for survival (Braaten et al. 2008).
Depending on water velocities, shovelnose sturgeon larvae may drift from 94 to 250 km
after hatching (Braaten et al. 2008). Dams on the Missouri river have fragmented habitat;
thus, reducing the amount of unimpeded river for drifting larvae (Braaten et al. 2008).
White sturgeon Acipenser transmontanus larvae require extended drift downstream of
spawning areas and are likely to exhibit little or no recruitment in reduced river reaches
terminating in reservoirs (Jager et al. 2002). Shovelnose sturgeon are thought to spawn
over rock or gravel substrate in tributaries or along main river channels (Moos 1978;
Keenlyne 1997). The location of shovelnose sturgeon spawning substrate in the Missouri
River could determine how much unimpeded riverine habitat larvae have for drifting.
The purpose of this study was to document spawning related movements of
shovelnose sturgeon through the annual spring pulse of the upper Missouri River, identify
possible spawning locations relative to available larval drift distance, and assess substrate
selection during spawning conditions. The specific objectives were to (1) compare the
movements of spawning shovelnose sturgeon between years with differing spring
hydrographs, (2) compare movements among different reproductive categories, (3)
document spawning locations of female shovelnose sturgeon relative to larval drift
distance, and (4) to document substrate availability in the upper Missouri River and
48
assess use relative to reproductive category. Based on what is currently known and
hypothesized regarding shovelnose sturgeon reproductive biology, I hypothesized that
shovelnose sturgeon would make upstream movements to spawn, movement of spawning
fish would be greater in a year with higher discharge, and that spawning fish would have
greater movements than reproductively inactive fish.
Study Area
The study area is located in north central Montana and includes the Missouri
River from a point approximately 10 km upstream of Fort Benton, MT (river kilometer
[rkm] 3,350, measuring from the confluence with the Mississippi River), to
approximately 40 km upstream of the headwaters of Fort Peck Reservoir (rkm 3,040)
(Figure 3.1). This section of the Missouri River was selected because it represents the
majority of main stem shovelnose sturgeon habitat between Morony Dam (rkm 3,388)
and the headwaters of Fort Peck Reservoir (rkm 3,000). Several dams exist in the
Missouri River system upstream of the study area, with three dams operated by the U.S.
Bureau of Reclamation (USBR) having the most effect on riverine flow and habitat
(Gardner and Jensen 2011). These include Canyon Ferry Dam on the Missouri River
near Helena, MT (rkm 3,626), Tiber Dam, on the Marias River near Chester, MT (rkm
129, measuring from the confluence with the Missouri River), and Gibson Dam (rkm
163, measured from confluence with the Missouri River) on the Sun River (Gardner and
Jensen 2011). Operation of these dams has altered the temperature and sediment
transport regimes and moderated the peaks and troughs of the average hydrograph, but
49
timing of peak discharge is similar to average conditions (Bellgraph et al. 2008).
Although the hydrograph is affected by these dams, the Missouri River above Fort Peck
Reservoir is considered the least hydrologically altered section of the Missouri River
(Galat and Lipkin 2000; Pegg et al. 2003; Galat et al. 2005). Further, the Missouri River
above Fort Peck Reservoir has many of the natural features common to unregulated rivers
including islands, alluvial bars, secondary channels, and backwaters. A combination of
factors contribute to the retention of natural features in this river section including limited
reservoir storage capacity, multi-use management of reservoirs, and numerous
unregulated tributaries that limit the influence of upstream impoundments (Scott et al.
1997; Galat and Lipkin 2000; Pegg et al. 2003).
Methods
Fish Capture and Surgery
Male shovelnose sturgeon, gravid female shovelnose sturgeon (stage IV;
Wildhaber et al. 2007), and reproductively inactive female shovelnose sturgeon (≤ stage
III; Wildhaber et al. 2007) were captured and implanted with radio transmitters at three
locations in the upper Missouri River in 2008 and 2009 (Table 3.1; Figure 3.1). The
three sampling locations were chosen such that individuals from the upper, middle, and
lower sections of the study area were represented. Shovelnose sturgeon were captured
from 23 April to 15 May in 2008, and from 13 to 21 April in 2009 using drifting trammel
nets (1.8 m deep x 45.8 m long, 2.5 cm mesh inner panel, 25.4-cm mesh outer panels).
Prior to transmitter implantation, gonads were visually inspected to determine sex and
reproductive stage (DeLonay et al. 2007). Shovelnose sturgeon were selected for
50
transmitter implantation if they were ≥ 1.9 kg and presumed to be either a male, gravid
female, or reproductively inactive female after macroscopic inspection of the gonad.
Transmitters were 16 x 73 mm, weighed 26 g, and had an estimated battery life of 1624 d
(Lotek Wireless, Inc., Newmarket, Ontario, Canada). Transmitters were selected to be as
light as possible while maximizing battery life and did not exceed 2% of fish body weight
(Winter 1996). Transmitter-implantation surgery was performed using methods modified
from Ross and Kleiner (1982) and Summerfelt and Smith (1990). All fish were placed
into a holding tank for 20 minutes to allow for recovery from the surgery, and were
released near the capture location. An effort was made to recapture previously classified
gravid females to determine spawning success during July-August in 2008 and 2009;
recapture was attempted with targeted drifting trammel nets after a fish was located via
telemetry (see methods below).
Reproductive Assessment
Ovarian tissue samples were collected from all females at the time of transmitter
implantation to corroborate reproductive stage classifications made in the field, and
samples were also collected from previously classified gravid females recaptured after
the spawning period to determine spawning success. Ovarian tissue samples were
preserved in 10% buffered formalin in the field and returned to the laboratory where they
were embedded in paraffin, sectioned at 5 µm, and stained by periodic acid Schiff stain
(Luna 1968). Slides were examined under a compound scope (Olympus America Inc.,
model BX41 40x-400x), and the germ cells were scored for stage of maturation according
to the protocol of Wildhaber et al. (2007).
51
Females were segregated into four categories based on stage of maturation and
condition of the gonad: confirmed spawners, potential spawners, atretic, or
reproductively inactive. Indications of successful spawning include the presence of postovulatory follicles and an absence of atretic bodies. If post-ovulatory follicles were
found in an ovarian tissue sample from a previously classified gravid female that was
recaptured post-spawn, the fish was classified as a confirmed spawner (Crim and Glebe
1990; Wildhaber et al. 2007). Previously classified gravid females that were not
recaptured post-spawn were classified as potential spawners. If no post-ovulatory
follicles were present and a large number of atretic bodies were observed the fish was
classified as atretic. Female shovelnose sturgeon require multiple seasons between
spawns for gonadal development (Billard and Lecointre 2001; Colombo et al. 2007;
DeLonay et al. 2009; Tripp et al. 2009); thus, confirmed spawning female shovelnose
sturgeon captured in July-August of 2008 were considered reproductively inactive in
2009.
Hydrographs and Water Temperature Profiles
To compare sturgeon movements through the course of the spring pulse, the
hydrograph was partitioned into three phases based on the putative reproductive lifehistory requirements of shovelnose sturgeon (Figure 3.2). The pre-suitable spawning
phase for each year was defined as the period between 1 May (arbitrarily selected based
on when sampling started) until the peak of the hydrograph, which was on 28 May in
2008 and on 3 June in 2009. The suitable spawning phase was defined as the descending
limb of the hydrograph, coincident to temperatures above 12° C, the lower threshold for
52
spawning and embryonic survival, until water temperature in the river exceeded 24° C,
the upper threshold of temperatures suitable for spawning and embryonic survival (K.
Kappenman and M. Webb, U.S. Fish and Wildlife Service, unpublished data). The
discharge thresholds were based on shovelnose sturgeon spawning in the upper Missouri
River that was documented to occur on the descending limb of the hydrograph,
coincident to temperatures within the suitable ranges for spawning and embryonic
survival (Goodman 2009). In 2008, suitable spawning conditions were from 29 May to
26 July, and in 2009 from 4 June to 22 July. The post-suitable spawning phase was
defined as the period after water temperatures in the river exceeded 24° C until the end of
tracking (31 August). Hydrographs were derived by averaging the mean daily discharge
recorded at three USGS gauging stations on the Missouri River within the study site: Fort
Benton (rkm 3,337), Virgelle Ferry (rkm 3,274), and Landusky (rkm 3,090). For
comparison to average discharge, data from the same three USGS gauging stations were
used to calculate the mean daily discharge averaged over the years 1935 to 2005. Water
temperature data were derived by averaging the mean daily water temperature (°C) from
two continuous-reading temperature loggers at rkm 3,092 and rkm 3,304 in 2008, and at
rkm 3,092 and rkm 3,193 in 2009.
Tracking
Individual fish were tracked weekly from May through August in 2008 and 2009
to estimate the direction and extent of movements associated with the pre-suitable
spawning, suitable spawning, and post-suitable spawning phases of the hydrograph.
Telemetered fish were located using a portable SRX-400 Lotek receiver (Lotek Wireless,
53
Inc., Newmarket, Ontario, Canada) equipped with a boat-mounted, four-element Yagi
antenna. More precise locations were determined by triangulation or by passing the boat
directly over the fish using a three-element hand held Yagi antenna (Winter 1996;
Blanchfield et al. 2005). This method has been shown to be accurate to within five
meters by means of blind tests using transmitters placed in the river (Gerrity et al. 2008;
Bellgraph et al. 2008). Latitude, longitude, and river kilometer were measured at each
fish location using a boat mounted GPS unit. Data from eight autonomous land-based,
continuous recording radio-receiving stations (Figure 3.1) were also incorporated into the
data set. These stations were located on the bank adjacent to the river and each station
consisted of a continuously recording, solar powered SRX-400 Lotek receiver attached to
two four-element Yagi antenna.
Movement and Movement Rates
Mean daily total and net movement rates were calculated for each fish for each
phase of the hydrograph. Total daily movement rate was calculated for a given fish by
dividing the distance in river kilometers between successive relocations by the number of
days elapsed between relocations (White and Garrott 1990). Net daily movement rate
was calculated by dividing the difference in river kilometer between successive
relocations by the number of days that elapsed between relocations such that a positive
rate indicated upstream movement, a negative rate indicated downstream movement, and
a rate of zero indicated no movement (Bramblett 1996). Additional movement likely
occurs between relocations; thus, all movement data represent a minimum for the period
between relocations (Rogers and White 2007). To identify possible spawning locations,
54
shovelnose sturgeon movements were plotted by category through the hydrograph phases
and visually assessed. In addition, mean river kilometer for locations of individual,
successfully spawning females during suitable spawning conditions were plotted and
visually assessed.
Substrate
In 2009 during the suitable spawning phase the predominant substrate at
shovelnose sturgeon relocations was determined by “feeling” the bottom of the river
channel using steel conduit (Bramblett and White 2001; Bellgraph et al. 2008; Gerrity et
al. 2008). Substrate within the study area was classified as: fines (0-4 mm in diameter),
gravel (5-64 mm), and cobble (65-300 mm). Blind tests in areas with known substrate
validated this method and substrate samples were obtained in these areas to validate size
classifications. The availability of substrate classes was estimated by dividing the study
area by river kilometer and randomly selecting two substrate sampling locations per river
kilometer (N = 558).
Data Analysis
Individual radio-tagged fish were the experimental unit for all applicable
statistical tests, and normality for all variables was assessed using residual and normal
probability plots. To reduce the chance of making a type II error, an α = 0.1 was
established a priori for all analyses. Discharge, water temperature, and rates of change
for each hydrograph phase were compared between years using a two-sample t-test after
data were log10 transformed to correct for non-normal distribution. Repeated measures
55
(with individual fish as the repeated variable) analysis of variance (ANOVA) was used to
test the hypotheses that there were no differences in total and net movement rates
between confirmed spawning females in 2008 and 2009. If movement rates did not differ
significantly, years were pooled to increase statistical power and repeated measures
ANOVA was used to test the hypotheses that there were no differences in total and net
movement rates among hydrograph phases.
Repeated measures ANOVA was also used to test the hypotheses that there were
no differences in total and net movement rates among reproductive categories (i.e., males,
potentially spawning females, confirmed spawning females, atretic females, and
reproductively inactive females) within each year. If movement rates did not differ
significantly, reproductive categories were pooled to increase statistical power and
repeated measures ANOVA was used to test the hypotheses that there were no
differences in total and net movement rates among hydrograph phases for each year.
Bonferroni multiple-comparisons test was used to control the experimentwise error rate
for all repeated measures ANOVAs (Rogers and White 2007). Sample sizes within
reproductive categories were relatively small; however, the repeated measures ANOVA
used to analyze the data takes into consideration the variation within individuals, thus
offering high statistical power relative to sample size (Ott 1993; Kutner et al. 2004).
One-way chi-square log-likelihood tests were conducted to test the null
hypotheses that substrate use within each substrate category was in proportion to
availability (Manly et al. 2002). Although some expected values used for chi-square
56
analyses were less than the recommended minimum of five, chi-square tests are robust to
smaller expected values (Roscoe and Byars 1971, Lawal and Upton 1984, Ott 1993).
Results
Hydrographs and Water Temperature Profiles
Daily discharge during the pre-suitable spawning phase was significantly higher
in 2008 than in 2009 (t = -6.54, P < 0.001), and peak discharge was 45 % higher in 2008
than in 2009 (Figure 3.2; Table 3.2). Daily discharge was also significantly higher during
the suitable spawning phase in 2008 than in 2009 (t = 7.18, P < 0.001), but daily
discharge did not differ significantly between years during the post-suitable spawning
phase (t = -0.11, P = 0.91). Rate of change in daily discharge differed between years for
the pre-suitable spawning phase (t = 1.84, P = 0.07), and was significantly higher in
2008. Rate of change in daily discharge did not differ significantly between years for the
suitable spawning phase (t = -1.14, P = 0.26), or the post-suitable spawning phase (t =
0.27, P = 0.78) (Figure 3.2; Table 3.2). In addition to higher peak discharge achieved at a
higher rate, discharge levels remain high for a greater duration in 2008 than in 2009. For
example, in 2008 discharge remained above the highest rate observed in 2009 (578 m3/s)
for 28 days, while discharge in 2009 began to decrease sharply immediately after the
peak was achieved (Figure 3.2). Discharge in 2008 was higher than average discharge
during the pre-suitable and suitable spawning phases, while discharge in 2009 during
these hydrograph phases more closely approximated average values. However, the
duration of increased discharge in 2008 more closely resembled that of the average
57
discharge, while the peak in discharge in 2009 was of a shorter duration. During the
post-suitable spawning phases discharge in both years was similar to the average
discharge values (Figure 3.2).
Daily water temperature did not differ significantly between years for the presuitable spawning phase (t = 0.14, P = 0.89) or the post-suitable spawning phase (t = 1.56, P = 0.62). However, daily water temperature differed significantly between years
during the suitable spawning phase (t = -2.33, P = 0.02), and was lower for a longer
period in 2008. The difference observed in water temperature between years during the
suitable spawning phase is likely a function of the prolonged period of high discharge in
2008 (Figure 3.2). Rate of change in daily water temperature did not differ significantly
between years for the pre-suitable spawning phase (t = -1.56, P = 0.13), suitable
spawning phase (t = 0.43, P = 0.66), and post-suitable spawning phase (t = 0.05, P =
0.96) (Figure 3.2; Table 3.2).
Movement and Movement Rates
Despite the higher, more prolonged discharge and the lower water temperature in
2008, mean net movement rate (F1, 17 = 0.24, P = 0.63) and mean total movement rate
(F1, 17 = 1.69, P = 0.21) of confirmed spawning female shovelnose sturgeon did not differ
significantly between years within hydrograph phases (Table 3.3). When years were
pooled, mean net movement rate (F2, 36 = 2.99, P = 0.06) and mean total movement rate
(F2, 36 = 9.49, P ˂ 0.001) of confirmed spawning female shovelnose sturgeon differed
significantly among hydrograph phases. Net movement rate of confirmed spawning
females was negative during the pre-suitable spawning phase indicating overall
58
downstream movement, while upstream movement was highest during the post-suitable
spawning phase movement. Net movement rate during the suitable spawning phase was
near zero, indicating nearly equal upstream and downstream movement (Table 3.3).
Interestingly, mean total movement rate was highest during the suitable spawning phase,
indicating that while upstream and downstream movement was nearly equal, confirmed
spawning females were moving the most during this hydrograph phase (Table 3.3). The
greatest upstream and downstream weekly mean net movement rates of confirmed
spawning females during both years were observed when water temperature was between
18 and 22 °C and discharge was 200-500 m3/s; interestingly at temperatures between 12
and 14 °C, net movement rates were similar at low (100-300 m3/s) and high (700-800
m3/s) discharge (Figure 3.3). The highest weekly mean total movement rates during both
years were at temperatures between 18 and 22 °C, when discharge was 700-800 m3/s in
2008 and 300-600 m3/s in 2009 (Figure 3.4). At temperatures below 18 °C and above 22
°C, weekly total movement rates for both years decreased at all discharges (Figure 3.4).
In 2008, mean net movement rate (F3, 27 = 0.03, P = 0.99) and mean total
movement rate (F3, 27 = 0.94, P = 0.44) did not differ significantly among reproductive
categories within hydrograph phases (Table 3.4). However, both net movement rate (F2,
58
= 3.58, P = 0.03) and total movement rate (F2, 58 = 14.12, P ˂ 0.001) differed among
hydrograph phases when reproductive categories were pooled (Table 3.4). In 2008, fish
in all reproductive categories made upstream and downstream movements during the
suitable spawning phase (Figure 3.5; Table 3.4). Not only were the lowest negative (1.83 km/d, overall downstream movement) and highest positive (1.91 km/d, overall
59
upstream movement) weekly mean net movement rates during the suitable spawning
phase, but variation around weekly mean net movement rates was also highest (Figure
3.5). For example, the mean of 90% confidence intervals for weekly mean net movement
rates for all reproductive categories pooled was highest during the suitable spawning
phase (mean = 2.31 km/d; SD = 1.49), compared to the pre-suitable (mean = 1.41 km/d;
SD = 0.77), and post-suitable (mean = 0.66 km/d; SD = 0.30) spawning phases. Overall,
total movement rates were highest during the suitable spawning phase at 1.63 km/d,
compared to 0.91 km/d during the pre-suitable spawning phase, and 0.47 km/d during the
post-suitable spawning phase (Table 3.4). This corresponds with the high variation
observed in weekly mean net movement rates during the suitable spawning phase as fish
increased movement upstream and downstream (Figure 3.5). Although no significant
differences were observed in mean total movement rates among reproductive categories,
the highest weekly mean total movement rates of males occurred 3-4 weeks before the
highest weekly mean total movement rates of females (Figure 3.6).
In 2009 mean net movement rate data were similar to those in 2008, with net
movement rates having no significant differences among reproductive categories within
hydrograph phases (F4, 51 = 0.65, P = 0.63), but differing among hydrograph phases when
reproductive categories were pooled (F2, 107 = 3.51, P = 0.03) (Table 3.5). As in the
previous year, fish in all reproductive categories made upstream and downstream
movements during the suitable spawning phase (Figure 3.7; Table 3.5). The highest
upstream weekly mean net movement rate in 2009 was during the suitable spawning
phase (2.29 km/d, overall upstream movement); however the highest downstream weekly
60
mean net movement rate was observed in males during the pre-suitable spawning phase (1.49 km/d, overall downstream movement) (Figure 3.7). As in 2008, the mean of 90%
confidence intervals for weekly mean net movement rates for all reproductive categories
pooled was highest during the suitable spawning phase (mean = 2.06 km/d; SD = 1.63),
compared to the pre-suitable (mean = 1.35 km/d; SD = 1.30), and post-suitable (mean =
0.99 km/d; SD = 0.56) spawning phases (Figure 3.7). While no significant differences
were observed in mean net movement rates among reproductive categories within
hydrograph phases in 2009, weekly mean net movement rates of reproductively inactive
females did not exhibit the variation observed in the other categories (Figure 3.7).
Weekly mean net movement rates of reproductively inactive females were centered on
zero for all hydrograph phases, and had little variation indicating a lack of movement
upstream or downstream. For all hydrograph phases, the highest positive weekly mean
net movement rate of reproductively inactive females was 0.25 km/d (overall upstream
movement) and the lowest negative weekly mean net movement rate was -0.37 km/d
(overall downstream movement), and the mean of 90% confidence intervals for weekly
mean net movement rates was low (mean = 0.41 km/d; SD = 0.14) (Figure 3.7). Unlike
2008, mean total movement rates in 2009 differed among reproductive categories within
hydrograph phases (F4, 51 = 3.76, P = 0.01) with reproductively inactive females moving
significantly less than other reproductive categories (Table 3.5). As in the previous year,
weekly mean total movement rates of males in 2009 peaked earlier than females, with the
highest total movement rates of males occurring 4-5 weeks prior to the highest female
total movement rates (Figure 3.8).
61
Shovelnose sturgeon migration patterns were highly variable with respect to
capture location but similar between years (Figures 3.9 and 3.10). For example,
shovelnose sturgeon initially located near Coal Banks did not exhibit extensive upstream
migrations while those initially located near Fred Robinson Bridge did not migrate
downstream (Figures 3.9 and 3.10). Migration patterns within reproductive categories
were similar between years; for example, in both years many of the males initially
located at Coal Banks (rkm 3,267) did not migrate, with only one male migrating
downstream. In contrast, in both years most males initially located near Judith Landing
(rkm 3,193) either migrated upstream or downstream, with only one male not migrating
(Figures 3.9 and 3.10). Migrations of confirmed spawning females initially located near
Coal Banks were similar between years with some fish not migrating while others
migrated downstream (Figures 3.9 and 3.10). Confirmed spawning females initially
located near Judith Landing in 2008 migrated upstream or downstream, while confirmed
spawning females in 2009 did not migrate or migrated downstream. Most confirmed
spawning females initially located near Fred Robinson Bridge in both years made no
migration, with one fish migrating upstream (Figures 3.9 and 3.10). Migration patterns
of potentially spawning females initially located near Coal Banks and Judith Landing
were similar to those of confirmed spawners from the same areas. However in both
years, potentially spawning females initially located near Fred Robinson Bridge made
extensive upstream migrations (Figures 3.9 and 3.10), indicating that some gravid
females from this area likely spawn at upstream locations. Atretic females exhibited
similar migrations to confirmed spawning females with one fish initially located near
62
Coal Banks migrating downstream, one initially located near Judith Landing migrating
upstream, and those initially located near Fred Robinson Bridge not migrating (Figures
3.9 and 3.10). Reproductively inactive females did not exhibit the extensive migrations
observed in the other reproductive categories (Figure 3.10). Confirmed spawning
females were clustered in two large areas during the suitable spawning phase; one
between rkm 3,250 and rkm 3,300, and the other between rkm 3,200 and rkm 3,075
(Figure 3.11).
Substrate
Cobble is the predominant substrate from rkm 3,340, upstream of Coal Banks, to
rkm 3,130, slightly upstream of Fred Robinson Bridge (Figure 3.12). The predominant
substrate below Fred Robinson Bridge is fines and sand (Figure 3.12). During the
suitable spawning phase, substrate use by fish within each reproductive category was not
in proportion to availability (Table 3.6; Figure 3.13). Fish in all reproductive categories
used gravel and fines at a higher proportion than were available, while use of cobble
varied with reproductive category (Figure 3.13). Mean percent of locations and
associated upper 90% confidence limits for all reproductive categories were higher than
the estimated availability of gravel and fines. Mean percent of locations for all
reproductive categories were lower than the estimated availability of cobble, indicating
that overall use of cobble was in lower proportion to estimated availability. However,
males, confirmed spawning females, and atretic females had upper 90% confidence limits
higher than the estimated availability of cobble substrate, while potentially spawning
females and reproductively inactive females had upper 90% confidence limits lower than
63
the estimated availability of cobble substrate. Care should be taken when interpreting
these confidence intervals as sample sizes for potentially spawning females (N = 17) and
reproductively inactive females (N = 17) were larger than those for males (N = 9),
confirmed spawning females (N = 10), and atretic females (N = 3).
Discussion
I rejected my hypothesis that shovelnose sturgeon in the upper Missouri River
would migrate upstream to spawn as I documented confirmed spawning female
shovelnose sturgeon not migrating or migrating downstream. Why all shovelnose
sturgeon did not migrate upstream to spawn in the upper Missouri River is unknown.
However, it is possible that fish resident to the furthest upstream extant of suitable
shovelnose sturgeon habitat near Coal Banks cannot make upstream migrations without
encountering unsuitable conditions. Conversely, fish resident to the area near Fred
Robinson Bridge cannot make extensive migrations downstream without encountering
the headwaters of Fort Peck Reservoir. The hypothesis that movement of spawning fish
would be greater in a year with higher discharge was rejected. Despite differences in
discharge and water temperature between years, movement rates and migration patterns
were similar. While females in this study were documented spawning successfully, the
ranges of discharge in this study were limited to those observed in 2008, an above
average discharge year, and 2009, which was more representative of average discharge.
How discharge outside of these ranges affects spawning migrations and how discharge
affects recruitment of shovelnose sturgeon in the upper Missouri River is unknown and
64
requires investigation. I failed to reject the hypothesis that spawning shovelnose sturgeon
would have greater movements than reproductively inactive females. Reproductively
inactive female shovelnose sturgeon did not exhibit the extensive migrations seen in other
reproductive categories and had significantly lower total movement rates. As the impetus
for migration is lacking (i.e., spawning) reproductively inactive females do not expend
energy migrating long distances. This understanding of the disparity in movements
between spawning and reproductively inactive fish may benefit future telemetry studies.
Total movement rates and the variation in net movement rates of confirmed
spawning, potentially spawning, and atretic females were highest during suitable
spawning conditions, and then declined during post-suitable spawning conditions to
similar values as observed in pre-suitable conditions. In both years, movements of males
were highest 3-5 weeks earlier than females. This is consistent with reported spawning
behavior of males in other sturgeon species; males arrive at spawning sites prior to
females, and stay longer as they attempt to maximize the opportunity for successful
mating (Fox et al. 2000; Paragamian and Kruse 2001; Bruch and Binkowski 2002). The
timing of increased movements in female shovelnose sturgeon in the upper Missouri
River coincides with the increase in the proportion of mature (stage IV) female
shovelnose sturgeon observed in the Mississippi River, which was concurrent with
increasing discharge and temperatures between 17-21° C (Tripp et al. 2009). The
increase in the proportion of mature females observed in the Mississippi River was
followed by an increase in the proportion of spent (stage VI) females approximately 1-2
months later (Tripp et al. 2009). Movements of atretic females in the upper Missouri
65
River were similar to confirmed and potentially spawning females indicating that while
failing to spawn, atretic females made similar spawning migrations. Similar patterns of
movement between successfully spawning female and atretic female shovelnose sturgeon
has been documented in a study in the Missouri River below Gavins Point Dam
(Wildhaber et al. 2011). My results corroborate that observation of a migration alone is
not sufficient evidence of successful spawning (Wildhaber et al. 2011). Female sturgeon
can fail to spawn and undergo follicular atresia when the appropriate cues for spawning
such as the presence of suitable habitat or mates are lacking (Auer 1996, 2004; Webb et
al. 1999, 2001; Linares-Casenave et al. 2002; Wildhaber et al. 2011); the conditions that
cause follicular atresia in shovelnose sturgeon in the upper Missouri River are unknown.
Reproductively inactive females did not make extensive spawning migrations in the
upper Missouri River. Reduced movements in reproductively inactive adult shovelnose
sturgeon compared to spawners were also observed in the Missouri River below Gavins
Point Dam (DeLonay et al. 2009).
Shovelnose sturgeon had variable migration patterns during the spawning period,
with some fish displaying no migration and others migrating upstream or downstream.
The migrations observed in the upper Missouri River do not support the hypothesis that
sturgeon species only make upstream migrations for spawning with downstream
migrations associated only with feeding (Bemis and Kynard 1997). This is also in
contrast to studies on the Missouri River below Gavins Point Dam that report gravid
female shovelnose sturgeon that migrate downstream or do not migrate fail to spawn,
with only females that migrate upstream spawning successfully (DeLonay et al. 2007,
66
2009). The distances travelled by confirmed spawning female shovelnose sturgeon in the
upper Missouri River (mean, 141.39 km; SD, 85.78) were also less than in shovelnose
sturgeon populations downstream (mean, 215.68 km; SD, 148.01; DeLonay et al. 2007).
It is possible that these discrepancies are due to the fragmented habitat in the upper
Missouri River. An estimated 310 km of contiguous, suitable habitat exists for
shovelnose sturgeon upstream from Fort Peck Reservoir, in contrast to 1,300 km of
unimpeded habitat available below Gavins Point Dam (DeLonay et al. 2007). Spawning
shovelnose sturgeon in the upper Missouri River cannot make extensive upstream
migrations without encountering naturally occurring unsuitable water temperatures and
reduced turbidity; conversely, fish attempting to migrate downstream encounter the
headwaters of Fort Peck Reservoir.
Shovelnose sturgeon spawned at multiple locations throughout the upper Missouri
River. Spawning at several locations over large patches of river has also been
documented in shovelnose sturgeon populations downstream in the Missouri River
(DeLonay et al. 2007; Delonay et al. 2009; Wildhaber et al. 2011). The highest densities
of confirmed spawning females during suitable spawning conditions were from rkm
3,125 to rkm 3,200. Confirmed spawning female shovelnose sturgeon did not appear to
use an approximately 50 km stretch of river between Coal Banks and Judith Landing.
Confirmed spawning female shovelnose sturgeon made migrations through this section of
river prior to suitable spawning conditions, but none remained there during suitable
spawning conditions. Why confirmed spawning female shovelnose sturgeon avoided this
area is unknown but a lack of spawning substrate can likely be ruled out, as the putative
67
spawning substrates of gravel and cobble are in abundance. A combination of habitat
factors is likely responsible for the lack of spawning in this area.
Shovelnose sturgeon larvae drift for 6 days post hatch (dph) and cover distances
from 94 to 250 km (Braaten et al. 2008). Using the minimum estimate of shovelnose
sturgeon larval drift distance, spawning would need to take place upstream of rkm 3,094
for larvae to avoid settling in the headwaters of Fort Peck Reservoir (rkm 3,000). In the
two years examined, 95%, of confirmed spawning females had mean locations during
suitable spawning conditions that were above this threshold. Not all confirmed spawning
female shovelnose sturgeon had mean locations above the minimum larval drift distance
threshold. However, it is likely that enough spawning is occurring upstream of this
threshold to maintain recruitment of shovelnose sturgeon.
This is the first study to quantify substrate distribution in the upper Missouri
River and to assess substrate use versus availability of fish of known reproductive
category during suitable spawning conditions. Shovelnose sturgeon have previously been
reported to use sand and gravel substrates throughout the Mississippi and Missouri river
systems (Becker et al. 1983; Carlson et al. 1985; Hurley et al. 1987; Curtis et al. 1997;
Quist et al. 1999; Bramblett and White 2001; Gerrity et al. 2008; DeLonay et al. 2009);
however, none of these studies took into account the sex or reproductive stage of fish.
Shovelnose sturgeon are thought to spawn over hard substrate such as rock, particularly
gravel (Keenlyne 1997), and other sturgeons spawn over gravel substrates (LaHaye et al.
1992; Paragamian et al. 2001, 2009; Manny and Kennedy 2002; Perrin et al. 2003).
While all reproductive categories used gravel and fines at a greater proportion than
68
availability, at the scale of this study and with the sample sizes used it is difficult to make
any inference as to substrate use of spawning shovelnose sturgeon, and more focused
study would further our understanding of spawning substrate requirements.
Recently, the hypothesis that discharge initiates spawning of shovelnose sturgeon
has been under increased scrutiny. Currently, it is thought that reproductive maturation
in shovelnose sturgeon is initiated many months prior to spawning with day length as the
environmental cue, while spawning is initiated by the proximate cue of water temperature
(DeLonay et al. 2009; Papoulias et al. 2011; Wildhaber et al. 2011). Scaphirhynchus spp.
were observed successfully spawning in the Mississippi River in both low and high water
years at suitable temperatures (Phelps et al. 2010), and multi-year studies on the lower
Missouri River have shown no discharge associated differences in physiological
measures of spawning readiness despite high variation in discharge between years and
study sections (DeLonay et al. 2009; Papoulias et al. 2011). My results support the
hypothesis that water temperature is a more likely proximate cue initiating spawning
migrations in shovelnose sturgeon than discharge. In 2008, a higher peak discharge was
achieved at a higher rate, and duration of high discharge was greater than in 2009.
Despite the differences in discharge, in both years female shovelnose sturgeon exhibited
similar movement rates and migrations while successfully spawning in the upper
Missouri River. Movements of confirmed spawning females were highest when water
temperatures were suitable for spawning regardless of discharge, indicating that
temperature may be a more important factor in the initiation of shovelnose sturgeon
spawning migrations. However, extended periods of high discharge have been shown to
69
result in protracted spawning and in turn a greater abundance of larval and age-0
Scaphirhynchus spp. (Goodman 2009; Phelps et al. 2010). How variations in discharge
affect the recruitment of shovelnose sturgeon in the upper Missouri River is unknown and
must be considered to effectively manage discharge for the benefit of shovelnose
sturgeon populations.
Acknowledgements
The U.S. Bureau of Reclamation and PPL Montana provided funding and support
for this research. I thank Robert Bramblett for manuscript reviews; Eli Cureton and Joel
Van Eenennaam for training and assistance with histological analyses; Eli McCord, Mike
Wente, Randy Rodencal, Ben Cox, Matt Jaeger, Michael Michelson, Nick Smith, Sue
Camp, Steve Leathe, Mike Meeuwig, Ben Goodman, Boone Richards, Justin Spinelli,
John Syslo, Peter Brown, Steven Ranney, and Mariah Talbott for their invaluable support
and assistance in the field, lab, and office. The Montana Cooperative Fisheries Research
Unit is jointly sponsored by the Montana Fish, Wildlife, and Parks, Montana State
University, and the U. S. Geological Survey.
70
Tables
Table 3.1. Sample size and tagging location of telemetered shovelnose sturgeon in the
Missouri River above Fort Peck Reservoir 2008-2009.
Location
Coal Banks (rkm 3,267)
Male
Gravid female
Reproductively inactive female
Year
2008 2009
5
7
Judith Landing (rkm 3,193)
Male
Gravid female
Reproductively inactive female
3
5
Fred Robinson Bridge (rkm 3,090)
Male
Gravid female
Reproductively inactive female
1
10
9
3
10
4
11
1
71
Table 3.2. Minimum, maximum, and mean (SD) daily discharge, daily water
temperature, and rates of change (ROC) for the Missouri River upstream of Fort Peck
Reservoir for all hydrograph phases 2008-2009. See text for hydrograph phase
definitions.
Variable
3
Discharge (m /s)
Discharge ROC (m3/s/d)
Year
Phase
Minimum
Maximum
Mean (SD)
2008
Pre-suitable
145
839
259 (192)
2009
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
206
156
299
215
161
823
207
578
530
216
542 (200)
185 (12)
440 (78)
313 (66)
185 (14)
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
-4
-65
-11
-15
-49
-18
165
45
11
39
36
18
26 (46)
-10 (20)
-1 (4)
8 (17)
-7 (17)
-1 (4)
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
10.26
11.33
20.88
6.58
13.54
17.88
17.59
23.25
24.12
18.39
23.67
25.37
13.28 (2.11)
18.21 (3.63)
22.50 (0.97)
13.46 (2.97)
19.54 (2.66)
21.32 (1.65)
Pre-suitable
Suitable
Post-suitable
Pre-suitable
-2.20
-1.61
-1.34
-1.18
1.82
1.38
0.60
1.51
0.02 (0.96)
0.20 (0.63)
-0.13 (0.55)
0.21 (0.63)
Suitable
Post-suitable
-1.33
-1.68
1.22
1.01
0.15 (0.68)
-0.09 (0.72)
2008
2009
Water temperature (° C)
2008
2009
Water temperature ROC (° C/d)
2008
2009
72
Table 3.3. Minimum, maximum, and mean ± 90% confidence interval (CI) net and total
movement rates (km/d) for confirmed spawning female shovelnose sturgeon in the
Missouri River upstream of Fort Peck Reservoir for all hydrograph phases 2008-2009.
Positive net movement rates represent upstream movement, negative rates represent
downstream movements. See text for hydrograph phase definitions.
Variable
Net movement rate
Year
2008
2009
Pooled
Total movement rate
2008
2009
Pooled
Phase
Minimum
Maximum
Mean ± 90% CI
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
-2.88
-2.03
-1.12
-2.24
-2.38
1.37
1.63
2.16
0.25
1.16
-0.09 ± 0.70
0.02 ± 0.54
0.29 ± 0.47
-0.69 ± 0.45
0.09 ± 0.54
Post-suitable
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
-1.16
-2.88
-2.38
-1.16
0.05
0.34
0.06
0.22
0.85
0.09
1.64
1.37
1.63
2.16
2.88
2.10
2.16
3.04
4.43
2.28
0.44 ± 0.43
-0.40 ± 0.41
0.06 ± 0.37
0.37 ± 0.31
0.86 ± 0.50
1.29 ± 0.36
0.55 ± 0.38
1.18 ± 0.53
1.92 ± 0.58
0.92 ± 0.45
Pre-suitable
0.05
3.04
1.03 ± 0.36
Suitable
0.34
4.43
1.62 ± 0.36
Post-suitable
0.06
2.28
0.75 ± 0.30
73
Table 3.4. Minimum, maximum, and mean ± 90% confidence interval (CI) net and total
movement rates (km/d) for reproductive categories of shovelnose sturgeon in the
Missouri River upstream of Fort Peck Reservoir for all hydrograph phases in 2008.
Positive net movement rates represent overall upstream movement, negative rates
represent downstream movement. Reproductive categories include males, confirmed
spawning females (CS), potentially spawning females (PS), and atretic females. See text
for hydrograph phase definitions.
Variable
Category
Net movement rate
Male
CS
PS
Atretic female
Pooled
Total movement rate
Male
CS
PS
Atretic female
Pooled
Phase
Minimum
Maximum
Mean ± 90% CI
Pre-suitable
Suitable
Post-suitable
-4.69
-0.16
-0.32
1.63
1.64
1.86
-0.78 ± 0.97
0.39 ± 0.36
0.41 ± 0.37
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
-2.88
-2.03
-1.12
-2.35
-2.88
-0.50
-0.33
-0.11
-0.04
0.07
1.12
0.50
1.37
1.63
2.16
0.86
3.34
1.69
0.30
0.10
0.09
4.69
1.17
0.60
-0.09 ± 0.70
0.02 ± 0.54
0.29 ± 0.47
-0.42 ± 0.44
0.39 ± 1.01
0.29 ± 0.37
0.01 ± 0.30
0.01 ± 0.10
0.02 ± 0.10
-0.39 ± 0.37
0.25 ± 0.37
0.31 ± 0.21
Pre-suitable
0.07
4.69
1.44 ± 0.86
Suitable
0.05
2.94
1.17 ± 0.61
Post-suitable
Pre-suitable
0.01
0.05
1.86
2.88
0.50 ± 0.33
0.86 ± 0.50
Suitable
0.34
2.10
1.29 ± 0.36
Post-suitable
0.06
2.16
0.55 ± 0.38
Pre-suitable
0.11
2.35
0.68 ± 0.33
Suitable
0.85
5.19
2.53 ± 0.79
Post-suitable
0.00
1.69
0.45 ± 0.31
Pre-suitable
0.07
0.36
0.26 ± 0.16
Suitable
0.07
1.64
1.00 ± 0.78
Post-suitable
0.04
0.09
0.06 ± 0.03
Pre-suitable
0.05
4.69
0.91 ± 0.32
Suitable
0.05
5.19
1.63 ± 0.37
Post-suitable
0.00
2.16
0.47 ± 0.18
74
Table 3.5. Minimum, maximum, and mean ± 90% confidence interval (CI) net and total
movement rates (km/d) for reproductive categories of shovelnose sturgeon in the
Missouri River upstream of Fort Peck Reservoir for all hydrograph phases in 2009.
Positive net movement rates represent overall upstream movement, negative rates
represent downstream movement. Reproductive categories include males, confirmed
spawning females (CS), potentially spawning females (PS), atretic females, and
reproductively inactive females (RI). See text for hydrograph phase definitions.
Variable
Category
Net movement rate
Male
CS
PS
Atretic female
Pooled
Total movement rate
Male
CS
PS
Atretic female
Pooled
Phase
Minimum
Maximum
Mean ± 90% CI
Pre-suitable
Suitable
Post-suitable
-4.69
-0.16
-0.32
1.63
1.64
1.86
-0.78 ± 0.97
0.39 ± 0.36
0.41 ± 0.37
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
Pre-suitable
Suitable
Post-suitable
-2.88
-2.03
-1.12
-2.35
-2.88
-0.50
-0.33
-0.11
-0.04
0.07
1.12
0.50
1.37
1.63
2.16
0.86
3.34
1.69
0.30
0.10
0.09
4.69
1.17
0.60
-0.09 ± 0.70
0.02 ± 0.54
0.29 ± 0.47
-0.42 ± 0.44
0.39 ± 1.01
0.29 ± 0.37
0.01 ± 0.30
0.01 ± 0.10
0.02 ± 0.10
-0.39 ± 0.37
0.25 ± 0.37
0.31 ± 0.21
Pre-suitable
0.07
4.69
1.44 ± 0.86
Suitable
0.05
2.94
1.17 ± 0.61
Post-suitable
Pre-suitable
0.01
0.05
1.86
2.88
0.50 ± 0.33
0.86 ± 0.50
Suitable
0.34
2.10
1.29 ± 0.36
Post-suitable
0.06
2.16
0.55 ± 0.38
Pre-suitable
0.11
2.35
0.68 ± 0.33
Suitable
0.85
5.19
2.53 ± 0.79
Post-suitable
0.00
1.69
0.45 ± 0.31
Pre-suitable
0.07
0.36
0.26 ± 0.16
Suitable
0.07
1.64
1.00 ± 0.78
Post-suitable
0.04
0.09
0.06 ± 0.03
Pre-suitable
0.05
4.69
0.91 ± 0.32
Suitable
0.05
5.19
1.63 ± 0.37
Post-suitable
0.00
2.16
0.47 ± 0.18
75
Table 3.6. Likelihood chi-square statistics for overall selection of substrate types by
shovelnose sturgeon reproductive category in the Missouri River above Fort Peck
Reservoir during the suitable spawning phase in 2009 (α = 0.1). Reproductive categories
include males, confirmed spawning females (CS), potentially spawning females (PS),
atretic females, and reproductively inactive females (RI); substrate types include cobble,
gravel, and fines.
Reproductive category
Males
CS
PS
Atretic females
RI
χ2
29.20
38.21
96.83
18.88
58.69
df
18
20
34
6
34
P
0.05
0.008
˂ 0.001
0.004
0.005
76
Figures
Figure 3.1. Map of Missouri and Marias rivers in Montana, with study area in detail.
77
900
Pre-suitable
Suitable spawning
30
Post-suitable
800
25
700
600
20
500
15
10
200
3
Mean daily discharge (m /s)
300
5
100
2008
0
May
0
Jun
Jul
Aug
Sep
Temperature
Discharge
Average discharge
900
Pre-suitable
Suitable spawning
Post-suitable
30
800
25
700
600
20
Mean daily water temperature (°C)
400
500
15
400
300
10
200
5
100
2009
0
May
0
Jun
Jul
Aug
Sep
Month
Figure 3.2. Hydrograph phases for the Missouri River upstream of Fort Peck Reservoir,
2008 and 2009 as defined by discharge and water temperature relative to shovelnose
sturgeon putative reproductive life history. Average discharge represents the mean daily
discharge averaged over 1935-2005. Hydrograph phases include pre-suitable, suitable,
and post-suitable spawning conditions (see text for hydrograph phase definitions).
78
2.0
1.5
1.0
-1.5
20
mean w
18
200
16
14
ater tem
perature
12
( oC)
100
Week
22
Weekly
ly mea
300
-2.0
harge
-1.0
n disc
-0.5
Weekly
mean n
800
700
600
500
400
0.0
(m 3/s)
0.5
e
ment
t move
/d)
rate (km
2008
2009
Figure 3.3. Weekly mean net movement rates of confirmed spawning female shovelnose
sturgeon in the Missouri River upstream of Fort Peck Reservoir 2008-2009 as a function of
weekly mean discharge and water temperature. Net movement rates above zero indicate
overall upstream movement, while rates below zero indicate overall downstream movement.
79
4.0
3.5
2.5
800
700
1.5
600
400
0.5
0.0
22
200
20
Weekly mean
18
16
water temperat
14
12
100
Weekly
300
mean d
500
1.0
ischarg
2.0
e (m 3/s
)
3.0
m
n total move
Weekly mea
ent rate (km/d
)
2008
2009
ure ( oC)
Figure 3.4. Weekly mean total movement rates of confirmed spawning female shovelnose
sturgeon in the Missouri River upstream of Fort Peck Reservoir 2008-2009 as a function of
weekly mean discharge and water temperature.
80
Pre-suitable
Suitable spawning
Post-suitable
Pre-suitable
Suitable spawning
Post-suitable
30
900
4
800
25
3
700
2
20
600
1
500
0
15
400
10
Net movement rate (km/d)
-2
-3
5
-4
-5
May
Male
Jun
Jul
Aug
CS
Sep
May
Jun
Jul
Aug
0
Sep
Temperature
Discharge
5
30
4
25
3
2
20
1
300
200
100
0
900
800
700
600
500
0
15
400
-1
10
-2
300
200
-3
5
100
-4
-5
May
Mean daily water temperature (°C)
-1
Mean daily discharge (m3/s)
5
PS
Jun
Jul
Aug
Atretic
Sep
May
Jun
Jul
Aug
0
0
Sep
Month
Figure 3.5. Mean net movement rates for male, confirmed spawning female (CS),
potentially spawning female (PS), and atretic female shovelnose sturgeon by hydrograph
phase in 2008. Symbols represent weekly mean net movement rates and error bars
represent 90% confidence intervals. Values greater than zero indicate movement
upstream, while values less than zero represent movement downstream.
81
9
Pre-suitable
Suitable spawning
Post-suitable
Pre-suitable
Suitable spawning
30
Post-suitable
900
8
800
25
7
700
6
20
600
5
500
15
10
Male
2
CS
5
1
0
May
0
Jun
Jul
Aug
Sep
May
Jun
Jul
Aug
Sep
Temperature
Discharge
9
30
8
25
7
6
20
5
300
200
100
0
900
800
Mean daily discharge (m3/s)
400
3
Mean daily water temperature (°C)
Total movement rate (km/d)
4
700
600
500
15
4
400
3
10
PS
2
Atretic
300
200
5
1
100
0
May
0
Jun
Jul
Aug
Sep
May
Jun
Jul
Aug
0
Sep
Month
Figure 3.6. Mean total movement rates for male, confirmed spawning female (CS),
potentially spawning female (PS), and atretic female shovelnose sturgeon by hydrograph
phase in 2008. Symbols represent weekly mean total movement rates and error bars
represent 90% confidence intervals.
82
Pre-suitable
Suitable spawning
Post-suitable
Pre-suitable
Suitable spawning
Post-suitable
30
900
4
800
25
3
700
2
20
600
1
500
0
15
10
-2
-3
5
-4
-5
May
Male
Jun
Jul
Aug
CS
Sep
May
Jun
Jul
Aug
Sep
5
Net movement rate (km/d)
0
30
4
25
3
2
20
1
Mean daily water temperature (°C)
400
-1
200
100
0
900
800
700
600
500
0
15
400
-1
10
-2
300
200
-3
5
100
-4
-5
May
300
Mean daily discharge (m3/s)
5
PS
Jun
Jul
Aug
Atretic
Sep
May
Jun
Jul
Aug
0
0
Sep
30
4
25
3
2
20
1
0
15
-1
10
-2
-3
5
-4
-5
May
RI
Jun
Jul
Aug
0
900
800
700
600
500
400
300
200
100
Mean daily discharge (m3/s)
5
Mean daily water temperature (°C)
Month
Temperature
Discharge
0
Sep
Month
Figure 3.7. Mean net movement rates for male, confirmed spawning female (CS),
potentially spawning female (PS), atretic female, and reproductively inactive female (RI)
shovelnose sturgeon by hydrograph phase in 2009. Symbols represent weekly mean net
movement rates and error bars represent 90% confidence intervals. Values greater than
zero indicate movement upstream, while values less than zero represent movement
downstream.
83
7
Pre-suitable
Suitable spawning
Post-suitable
Pre-suitable
Suitable spawning
Post-suitable
30
900
800
6
25
700
5
20
600
4
500
15
CS
Total movement rate (km/d)
2
10
1
5
0
May
0
Jun
Jul
Aug
Sep
May
Jun
Jul
Aug
Sep
7
30
6
25
5
20
4
300
200
100
0
900
800
700
Mean daily discharge (m3/s)
Male
Mean daily water temperature (°C)
400
3
600
500
15
400
3
PS
Atretic
2
10
300
200
5
1
0
May
100
0
Jun
Jul
Aug
Sep
May
Jun
Jul
Aug
0
Sep
30
6
25
5
20
4
15
3
RI
2
10
1
5
0
May
0
Jun
Jul
Aug
900
800
700
600
500
400
300
200
100
Mean daily discharge (m3/s)
7
Mean daily water temperature (°C)
Month
Temperature
Discharge
0
Sep
Month
Figure 3.8. Mean total movement rates for male, confirmed spawning female (CS),
potentially spawning female (PS), atretic female, and reproductively inactive female (RI)
shovelnose sturgeon by hydrograph phase in 2009. Symbols represent weekly mean total
movement rates and error bars represent 90% confidence intervals.
84
3350
Pre-suitable
Suitable spawning
Post-suitable
Pre-suitable
Suitable spawning
Post-suitable
3300
3250
3200
3150
River kilometer
3100
Male
3050
May
Jun
Jul
Aug
CS
Sep
May
Jun
Jul
Aug
Sep
3350
3300
3250
3200
3150
3100
PS
3050
May
Jun
Jul
Aug
Atretic
Sep
May
Jun
Jul
Aug
Sep
Month
Figure 3.9. Movement patterns of male, confirmed spawning female (CS), potentially
spawning female (PS), and atretic female shovelnose sturgeon in the Missouri River
above Fort Peck Reservoir, 2008. River kilometer designates distance upstream from the
confluence with the Mississippi River (rkm 0). In all panels, dotted lines represent fish
initially captured at Coal Banks (rkm 3,267), solid lines represent fish initially captured at
Judith Landing (rkm 3,193), and dashed lines represent fish initially captured at Fred
Robinson Bridge (rkm 3,090).
85
3350
Pre-suitable
Suitable spawning
Post-suitable
Pre-suitable
Suitable spawning
Post-suitable
3300
3250
3200
3150
3100
Male
3050
CS
3350
River kilometer
3300
3250
3200
3150
3100
PS
3050
Atretic
May
Jun
Jul
Aug
Sep
Month
3350
3300
3250
3200
3150
3100
3050
May
RI
Jun
Jul
Aug
Sep
Month
Figure 3.10. Movement patterns of male, confirmed spawning female (CS), potentially
spawning female (PS), atretic, and reproductively inactive female (RI) shovelnose
sturgeon in the Missouri River above Fort Peck Reservoir, 2009. River kilometer
designates distance upstream from the confluence with the Mississippi River (rkm 0). In
all panels, dotted lines represent fish initially captured at Coal Banks (rkm 3,267), solid
lines represent fish initially captured at Judith Landing (rkm 3,193), and dashed lines
represent fish initially captured at Fred Robinson Bridge (rkm 3,090).
86
3350
Coal Banks
Judith Landing
Fred Robinson Bridge
River kilometer
3300
3250
3200
3150
3100
2008
2009
3050
766
767
782
842
843
977
981
982
990
999 9111 9114 9116 9117 9120 9122 9123 9124 9125
Confirmed spawning female shovelnose sturgeon
Figure 3.11. Mean river kilometer of confirmed spawning female shovelnose sturgeon
during suitable spawning conditions in the Missouri river above Fort Peck reservoir, 2008
and 2009. Error bars represent 90% confidence intervals and in both panels triangles
represent fish initially captured at Coal Banks (rkm 3,267), squares represent fish initially
captured at Judith Landing (rkm 3,193), and circles represent fish initially captured at
Fred Robinson bridge (rkm 3,090). River kilometer designates distance upstream from
the confluence with the Mississippi River (rkm 0).
87
FRB
Judith Landing
Coal Banks
100
Percent
80
60
Cobble
Gravel
Fines/sand
40
20
0
3060
3100
3140
3180
3220
3260
3300
3340
River Kilometer
Figure 3.12. Distribution of substrate types in the Missouri River upstream of Fort Peck
Reservoir, 2009, as estimated from 558 randomly chosen points. Vertical lines indicate
locations of Fred Robinson Bridge (FRB), Judith Landing, and Coal Banks. River
kilometer designates distance upstream from the confluence with the Mississippi River
(rkm 0).
88
100
Male
CS
PS
Atretic
RI
Availability
Percent locations
80
60
40
20
0
Cobble
Gravel
Fines
Substrate
Figure 3.13. Substrate use by shovelnose sturgeon reproductive category versus
availability in the Missouri River upstream of Fort Peck Reservoir during the suitable
spawning phase, 2009. Substrate availability was estimated from 558 randomly chosen
points; error bars represent 90% confidence intervals. Reproductive categories include
males, confirmed spawning females (CS), potentially spawning females (PS), atretic
females, and reproductively inactive females (RI).
89
References
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CHAPTER FOUR
CONCLUSIONS
The goal of this thesis was to investigate the effects of variations in spring
discharge on movements of Scaphirhynchus spp. in the upper Missouri River. A
secondary goal was to provide information on the distribution of Scaphirhynchus spp.
locations relative to larval drift distance. In chapter two I compared movements of
Scaphirhynchus spp. among four years with hydrographs representing a range of
conditions from below average to above average discharge, with varying discharge rates
of change, and differing timing and duration of peak discharge. In addition, I compared
movement patterns and distributions of Scaphirhynchus spp. locations in the upper
Missouri River. I rejected the hypotheses that movements of Scaphirhynchus spp. would
be greater in a year with higher discharge, the highest movement rates would be
coincident to rising discharge, and that movements would be upstream. I failed to reject
the hypothesis that adult pallid sturgeon would move farther than juvenile pallid sturgeon
and shovelnose sturgeon. In chapter three I compared movements of spawning
shovelnose sturgeon between years with differing spring hydrographs, compared
movements among different reproductive categories, documented potential spawning
locations of female shovelnose sturgeon relative to larval drift distance, and documented
substrate distribution and availability in the upper Missouri River and assessed use
relative to reproductive category. I rejected the hypotheses that shovelnose sturgeon
would make upstream migrations to spawn and that movement of spawning fish would be
97
greater in a year with higher discharge. I failed to reject the hypothesis that spawning
fish would have greater movements than reproductively inactive fish. Below, I discuss
major findings while integrating chapters two and three, and discuss implications for
river management relative to Scaphirhynchus spp. ecology.
Rising spring discharge has been hypothesized to be a cue initiating spring
migrations in Scaphirhynchus spp. (Becker 1983; Keenlyne and Jenkins 1993; Goodman
2009; Tripp et al. 2009). The lack of sufficient discharge to initiate spawning migrations
has been proposed as a factor limiting Scaphirhynchus spp. spawning success in the
Missouri River and has guided management actions towards increasing spring discharge
(USFWS 2000, 2003). However, the results of this study provide evidence against the
hypothesis that increased discharge is a proximate cue initiating spring migrations.
Scaphirhynchus spp. did not exhibit differential movement rates in 2006 through 2009,
despite four differing hydrographs representing a range of conditions from below average
to above average discharge, with varying discharge rates of change, and differing timing
and duration of peak discharge. Additionally, despite the annual differences in discharge
peak, rate of change, and duration, in 2008 and 2009 female shovelnose sturgeon
exhibited similar movement rates and migrations while successfully spawning in the
upper Missouri River. Similar results were observed in shovelnose sturgeon in the
Kansas River wherein movements were not related to discharge (Quist et al. 1999), and in
the Mississippi River where Scaphirhynchus spp. were observed spawning in high and
low water years (Phelps et al. 2010). Multi-year studies on the lower Missouri River
have also shown no discharge associated differences in physiological measures of
98
spawning readiness despite high variation in discharge between years and study sections
(DeLonay et al. 2009; Papoulias et al. 2011). Rather than discharge, it has been
hypothesized that reproductive maturation in Scaphirhynchus spp. is initiated many
months prior to spawning with day length as the environmental cue, while spawning is
initiated by the proximate cue of water temperature (DeLonay et al. 2009; Papoulias et al.
2011; Wildhaber et al. 2011). Movement rates of confirmed spawning female shovelnose
sturgeon in the upper Missouri River in 2008 and 2009 were higher when water
temperatures were suitable for spawning, regardless of discharge; thus, providing support
for the hypothesis that water temperature is a more likely proximate cue initiating
spawning migrations in Scaphirhynchus spp.
In chapter three, when sex and reproductive stage of shovelnose sturgeon were
known, movement rates were significantly highest during the suitable spawning period
which was defined as the descending limb of the hydrograph when water temperature
was within the ranges suitable for spawning. However, in chapter two, where sex and
reproductive stage of shovelnose sturgeon were unknown, no differences were found in
movement rates among the ascending limb, descending limb, or post-descending limb.
The lack of a significant difference in movement rates when sex and reproductive stage
likely occurs when highly variable telemetry data obtained from fish of varying
reproductive categories are pooled. When sex and reproductive stage were known,
movement rates of reproductively inactive female shovelnose sturgeon were significantly
lower, as reproductively inactive females failed to undertake the extensive migrations
seen in males, potentially spawning females, and confirmed spawning females. This was
99
also documented in the Missouri River below Gains Point Dam, where reproductively
inactive adult shovelnose sturgeon had reduced movements compared to spawning fish
(DeLonay et al. 2009). Pooling data from reproductively inactive females with data from
other reproductive categories would likely affect results and potentially lead to incorrect
inferences. This indicates the need for identification of sex and reproductive stage of
study subjects when conducting telemetry studies.
Sturgeon are hypothesized to move upstream as spring discharge rises (Hall et al.
1991; Bemis and Kynard 1997; Wilson and Mckinley 2004); however, the patterns of
movement observed in the upper Missouri River do not support this hypothesis.
Scaphirhynchus spp. were observed moving upstream, moving downstream, and not
moving. When sex and reproductive stage of fish were known, female shovelnose
sturgeon were observed making spawning migrations upstream and downstream as well
as not migrating, yet still spawning successfully. Studies on the Missouri River below
Gavins Point Dam report gravid female shovelnose sturgeon that migrate downstream or
do not migrate fail to spawn, with only females that migrate upstream spawning
successfully (DeLonay et al. 2007, 2009). Why all shovelnose sturgeon did not migrate
upstream to spawn in the upper Missouri River is unknown. However, fish resident to
the furthest upstream extant of suitable shovelnose sturgeon habitat near Coal Banks
cannot make upstream migrations without encountering unsuitable conditions;
conversely, fish resident to the area near Fred Robinson Bridge cannot make extensive
migrations downstream without encountering the headwaters of Fort Peck Reservoir.
100
Shovelnose sturgeon locations were distributed farther upstream than juvenile
pallid sturgeon or adult pallid sturgeon locations. White sturgeon populations in
fragmented river sections terminating in reservoirs have reduced recruitment (Jager
2002), and the extended larval drift in pallid sturgeon may me a mechanism contributing
to the lack of recruitment in the upper Missouri River (Braaten et al. 2008). Pallid
sturgeon larvae may drift for 245 to 530 km and shovelnose sturgeon larvae may drift for
94 to 250 km depending on discharge (Braaten et al. 2008). This may be critical for
pallid sturgeon recruitment because the majority of locations in the four years of this
study were in the 75 km directly upstream of the Fort Peck Reservoir headwaters, and the
farthest upstream location was only 250 km above the headwaters, which is the
approximate minimum drift distance required for larval pallid sturgeon. Conversely,
shovelnose sturgeon locations were distributed approximately 350 km upstream of the
headwaters, and when sex and reproductive stage were known, 95% of confirmed
spawning female shovelnose sturgeon had mean locations during suitable spawning
conditions that were above the minimum larval shovelnose sturgeon drift threshold. Not
all confirmed spawning female shovelnose sturgeon had mean locations above the
minimum larval drift distance threshold; however, it is likely that enough spawning is
occurring upstream of this threshold to maintain recruitment of shovelnose sturgeon,
while the extended larval drift observed in pallid sturgeon may be contributing to the lack
of recruitment observed in the upper Missouri River.
The distribution of substrate in the upper Missouri River may partially explain the
different distributions of pallid sturgeon and shovelnose sturgeon locations. In the lower
101
Yellowstone River and the Missouri River above Lake Sakakawea, pallid sturgeon used
areas of fines and sand substrate more than areas of gravel or cobble substrate, while
shovelnose sturgeon were found most often in areas of cobble and gravel substrate
(Bramblett and White 2001). Juvenile pallid sturgeon in the upper Missouri River were
also found predominantly in areas with silt and sand substrate, with shovelnose sturgeon
using gravel and cobble substrate to a greater extent than juvenile pallid sturgeon (Gerrity
et al. 2008). In the upper Missouri River, cobble is the predominant substrate from rkm
3,340, upstream of Coal Banks, to rkm 3,130, slightly upstream of Fred Robinson Bridge,
while the predominant substrate below Fred Robinson Bridge is fines and sand. The
majority of pallid sturgeon locations in the four years of the study were in the area
dominated by fines and sand below rkm 3,130, while shovelnose sturgeon locations were
distributed across the study area.
My findings have implications for river management and indicate the need for
further research on Scaphirhynchus spp. in the upper Missouri River. Unlike other
populations, shovelnose sturgeon in the upper Missouri River were observed making
downstream migrations or not migrating while successfully spawning. This is possibly
due to the constrained habitat in the upper Missouri River, which is bounded upstream by
a naturally occurring unsuitable water temperature and turbidity gradient and downstream
by Fort Peck Reservoir. Scaphirhynchus spp. in the upper Mississippi River, the upper
Missouri River, and the Yellowstone and Missouri rivers above Lake Sakakawea
generally avoided impounded areas (Curtis et al. 1997; Bramblett and White 2001;
Gerrity et al. 2008). Manipulation of the Fort Peck Reservoir operational pool level can
102
determine the amount of lotic habitat available for Scaphirhynchus spp. in the upper
Missouri River (Gerrity et al. 2008). Reservoir operational pool level may also have
implications for the length of free-flowing river available for larval drift. The majority of
pallid sturgeon locations in the upper Missouri River were in the area below rkm 3,130
where sand is the predominant substrate type. Maintaining a reservoir pool level less
than full pool increases the amount of Scaphirhynchus spp. habitat available and the
length of river available for larval drift. For example, Fort Peck Reservoir operational
pool levels in 2004 made available and additional 60 km of lotic habitat that is inundated
when the reservoir is at full operational pool level (Gerrity et al. 2008).
Rising spring discharge is hypothesized to cue spawning migrations in
Scaphirhynchus spp. (Becker 1983; Keenlyne and Jenkins 1993; USFWS 2003;
Goodman 2009; Tripp et al. 2009). However, despite many studies investigators have
failed to find a link between discharge and spawning in Scaphirhynchus spp. (Papoulias
et al. 2011). Multi-year studies on the lower Missouri River have shown no discharge
associated differences in physiological measures of spawning readiness despite high
variations in discharge (DeLonay et al. 2009; Papoulias 2011), and it has been
hypothesized that water temperature is a more likely proximate spawning cue. My
findings provide further evidence that water temperature is a more likely proximate cue
than discharge in initiating spawning migrations of Scaphirhynchus spp. Movements of
Scaphirhynchus spp. did not differ among years despite variation in hydrographs, and
movement rates of confirmed spawning female shovelnose sturgeon peaked when water
temperatures were between 18 and 22 °C regardless of discharge. Thus, rather than
103
managing rivers for high spring discharge (USFWS 2000; 2003), it may be more
important to provide discharge such that water temperatures are suitable for spawning.
Although discharge did not affect movements in the upper Missouri River, prolonged
discharge has resulted in protracted spawning and a greater abundance of larval and age-0
Scaphirhynchus spp. (Goodman 2009; Phelps et al. 2010). Thus, discharge may be an
important factor in determining Scaphirhynchus spp. recruitment. Further research
should focus on how discharge affects Scaphirhynchus spp. recruitment in the upper
Missouri River, and explore the causative mechanisms between water temperature and
initiation of spawning.
104
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Geological Survey Scientific Investigations Report 2009-5201, Reston, Virginia.
Gerrity, P. C., C. S. Guy, and W. M. Gardner. 2008. Habitat use of juvenile pallid
sturgeon and shovelnose sturgeon with implications for water level management
in a downstream reservoir. North American Journal of Fisheries Management
28:832-843.
Goodman, B. J. 2009. Ichthyoplankton density and shovelnose sturgeon spawning in
relation to varying discharge treatments. Master’s thesis. Montana State
University, Bozeman, Montana.
Hall, J. W., T. I. J. Smith, and S. D. Lamprecht. 1991. Movements and habitats of
shortnose sturgeon, Acipenser brevirostrum in the Savannah River. Copeia
1991:695-702.
105
Jager, H. I., W. Van Winkle, J. A. Challender, K. B. Lepla, P. Bates, and T. D. Counihan.
2002. A simulation study of factors controlling white sturgeon recruitment in the
Snake River. Pages 127-150 in W. Van Winkle, P. J. Andres, D. H. Secor, and D.
A. Dixon, editors. Biology, management and protection of North American
sturgeon. American Fisheries Society, Symposium 28, Bethesda, Maryland.
Keenlyne, K. D., and L. G. Jenkins. 1993. Age at sexual maturity of the pallid sturgeon.
Transactions of the American Fisheries Society. 122:393-396.
Papoulias, D. M., A. J. DeLonay, M. L. Wildhaber, and D.E. Tillit. 2011.
Characterization of environmental cues for initiation of reproductive cycling and
spawning in shovelnose sturgeon Scaphirhynchus platorynchus in the lower
Missouri river, USA. Journal of Applied Ichthyology 27:335-342.
Phelps, Q. E., S. A. Tripp, W. D. Hintz, J. E. Garvey, D. P Herzog, D. E. Ostendorf, J. W.
Ridings, J. W. Crites, and R. A. Hrabik. 2010. Water temperature and river stage
influence mortality and abundance of naturally occurring Mississippi River
Scaphirhynchus sturgeon. North American Journal of Fisheries Management
30:767-775.
Quist, M. C., J. S. Tillma, M. N. Burlingame, and C. S. Guy. 1999. Overwinter habitat
use of shovelnose sturgeon in the Kansas River. Transactions of the American
Fisheries Society 128:522-527.
Tripp, S. J., Q. E. Phelps, R. E. Colombo, J. E. Garvey, B. M. Burr, D. P. Herzog, and R.
A. Hrabik. 2009. Maturation and reproduction of shovelnose sturgeon in the
middle Mississippi River. North American Journal of Fisheries Management
29:730-738.
USFWS (United States Fish and Wildlife Service). 2000. Biological opinion on the
operation of the Missouri River main stem reservoir system, operation and
maintenance of the Missouri River bank stabilization and navigation project,
and operation of the Kansas River reservoir system. USFWS, Bismarck, North
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Wildhaber, M. L., A. J. DeLonay, D. M. Papouilas, D. L. Galat, R. B. Jacobsen, D. G.
Simpkins, P. J. Braaten, C. E. Korschgen, and M. J. Mac. 2011. Identifying
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pallid and shovelnose sturgeons. Journal of Applied Ichthyology 27:462-469.
106
Wilson, J. A., and R. S. McKinley. 2004. Distribution, habitat, and movements. Pages
40-72 in G. T. O. LeBreton, F. W. Beamish, and R. S. McKinley, editors.
Sturgeons and paddlefish of North America. Kluwer Academic Publishers,
Dordrecht, The Netherlands.
107
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115
APPENDIX A
SHOVELNOSE STURGEON REPRODUCTIVE PHYSIOLOGY DATA
116
Table A.1. Mean polarization index (PI) ± SD for female shovelnose sturgeon at initial capture, and hormone levels and weights
of male and female shovelnose sturgeon at the beginning and end of the spring migration in the upper Missouri River, 2008-2009.
Samples at non-detectable levels (< 0.625 ng ml-1) are indicated by ND; values in parentheses represent the percent change in
hormone levels and weight between capture and recapture date.
Recapture date
PI ± SD
8/13
7/31
8/6
8/13
8/13
Estradiol (ng ml-1)
Capture Recapture
Weight (kg)
Capture Recapture
2.94
17.09
220.89
10.84
6.53
4.47
37.56
37.19
70.47
16.49
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
2.04
1.86
2.72
2.04
3.67
1.96 (-4)
1.91 (3)
2.58 (-5)
2.12 (4)
3.58 (-2)
8/6
7/29
7/29
8/24
8/19
8/6
0.16 ± 0.02
0.19 ± 0.03
0.21 ± 0.04
0.19 ± 0.03
0.21 ± 0.02
0.28 ± 0.02
20.30
26.38
24.11
28.80
25.37
21.83
0.92 (-95)
11.56 (-56)
0.94 (-96)
1.79 (-94)
0.65 (-97)
12.99 (-40)
2.81
4.53
3.70
15.89
2.57
4.82
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
2.12
1.84 (-13)
2.88
2.15
2.65
3.93
2.74 (-5)
1.88 (-13)
2.17 (-18)
3.92 (-1)
7/30
7/30
7/29
7/31
7/30
8/6
8/7
7/6
8/7
0.16 ± 0.02
0.16 ± 0.02
0.25 ± 0.03
0.22 ± 0.03
0.18 ± 0.03
0.16 ± 0.01
0.18 ± 0.02
0.17 ± 0.03
0.16 ± 0.02
12.07
0.52 (-96)
ND
ND (-99)
ND (-99)
0.30 (-99)
1.43 (-96)
ND (-99)
2.62 (-92)
0.69 (-98)
2.43
ND (-99)
ND
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
2.04
2.55
2.13
2.85
1.78 (-13)
2.06 (-19)
1.81 (-15)
2.21 (-22)
2.31
2.54
2.63
3.35
1.93 (-16)
2.31 (-9)
2.22 (-16)
2.97 (-11)
11.71
44.65
43.03
36.35
23.09
31.07
27.99
3.47
6.82
3.47
8.19
7.42
7.80
1.45
121
Fish ID
Capture date
Males
835
5/7
971
5/15
975
5/15
976
5/7
980
4/26
Atretic females
779
4/29
978
4/29
995
5/14
998
4/14
9108
4/20
9121
4/17
Confirmed spawning females
766
4/29
767
4/29
782
4/29
842
4/23
843
5/14
977
4/26
981
4/26
999
4/14
9111
4/20
Testosterone (ng ml-1)
Capture
Recapture
117
9114
9116
9117
9120
9122
9123
9124
9125
4/13
4/16
4/16
4/17
4/17
4/17
4/21
4/21
8/17
8/18
8/19
8/20
7/30
8/21
8/3
8/8
0.13 ± 0.02
0.14 ± 0.02
0.16 ± 0.02
0.20 ± 0.02
0.16 ± 0.02
0.14 ± 0.02
0.18 ± 0.02
0.18 ± 0.02
25.96
37.39
38.68
25.73
20.78
45.68
40.70
47.57
2.92 (-89)
5.94 (-84)
5.69 (-85)
1.91 (-93)
0.80 (-96)
2.04 (-96)
1.19 (-97)
0.28 (-99)
4.55
5.89
7.00
10.21
12.51
3.71
3.02
9.04
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
ND (-99)
2.54
3.59
2.54
3.63
2.81
3.15
3.25
2.22 (-13)
3.37 (-6)
2.32 (-9)
3.18 (-12)
2.67
2.47 (-12)
2.87 (-9)
2.57 (-21)
122