AN ABSTRACT OF THE THESIS OF
Sierra Koch Lewis for the degree of Master of Science in Fisheries Science presented
on May 3, 2013.
Title: Reconnecting Aquatic Habitats: Validating Historical Habitat Use by
Anadromous Fishes using Telemetry and Stable Isotope Analysis above Barriers.
Abstract approved:
____________________________________________________________________
David L.G. Noakes
I conducted a study to identify potential spawning habitat for anadromous
salmonids above a 60-year-old hydropower dam in the headwaters of the North
Umpqua River in Oregon. Like many other historical salmonid-bearing rivers, little
documentation exists for anadromous fish presence above potential natural obstacles
upstream of Soda Springs Dam. My prediction was that if migratory salmonids are
allowed to move upstream of the dam using a new fish ladder they would utilize
available upstream spawning habitat. I captured, radio-tagged, and transplanted wild
adult summer steelhead, O. mykiss, above Soda Springs Dam and tracked daily
movements for 1 year to observe habitat preferences. I also investigated evidence of
historical anadromy throughout the North Umpqua River basin using stable isotope
analysis of salmon-derived nitrogen (15N) in foliar samples from Douglas fir trees, P.
menziesii, (>100 years old). I tested the hypothesis that I could identify
undocumented, historical salmon spawning reaches above natural and anthropogenic
obstacles based solely upon δ15N foliar deposition patterns.
While tracking radio-tagged steelhead, I documented holding locations, the
timing of spawning activities, and the outmigration of kelts. Tagged fish did not show
any extensive movement in the river or tributaries above the dam. Most of the tagged
adults showed incremental daily movements near the release site or downstream into
the hydropower reservoir but did not travel further than 1 kilometer upstream in the
main river channel. I recorded movements and localized activities that suggested some
of the fish spawned and subsequently moved downstream towards the Pacific Ocean.
My observations indicated that spawning behavior of tagged fish above Soda Springs
Dam was delayed several weeks relative to fish spawning below the dam. There was
no evidence from telemetry that any fish moved upstream in Fish Creek, the newly
accessible habitat of interest in this project. My data suggest that steelhead will use
restored habitat above dams, but that re-colonization activities are variable and may be
affected by altered flow regimes within the restored habitat upstream of the dam and
intra-specific density dependence.
My stable isotope data indicated that the foliar δ15N deposition patterns were
confounded with elevation. I documented potential “salmon-derived” false positives
on the Umpqua National Forest above impassable waterfalls. Overall I found that
foliar δ15N deposition patterns were highly variable and unexpectedly indicated a
statistically significant negative correlation of foliar δ15N values with historical salmon
presence. My linear mixed-effects modeling suggests that the presence of salmon is
the most important indicator of foliar δ15N values, rather than the proximity of a
sampled tree to stream flow above or below migratory barriers. I was not able to use
the mixed effects model to identify previously undocumented salmon spawning
habitat. My results from the foliar technique suggest that the method may not be
universally applicable as has been previously suggested in the literature.
© Copyright by Sierra Koch Lewis
May 3, 2013
All Rights Reserved
Reconnecting Aquatic Habitats: Validating Historical Habitat Use by
Anadromous Fishes using Telemetry and Stable Isotope Analysis above Barriers
by
Sierra Koch Lewis
A THESIS
submitted to
Oregon State University
in partial fulfillment of
the requirements for the
degree of
Master of Science
Presented May 3, 2013
Commencement June 2013
Master of Science thesis of Sierra Koch Lewis
presented on May 3, 2013.
APPROVED:
____________________________________________________________________
Major Professor, representing Fisheries Science
____________________________________________________________________
Head of the Department of Fisheries and Wildlife
____________________________________________________________________
Dean of the Graduate School
I understand that my thesis will become part of the permanent collection of
Oregon State University libraries. My signature below authorizes release of my thesis
to any reader upon request.
____________________________________________________________________
Sierra Koch Lewis, Author
ACKNOWLEDGEMENTS
I express my sincere appreciation to David Noakes, my major advisor, for his
unwavering support and willingness to let me work my way upstream on my own
terms. He provided unflagging support across both time and space whenever I asked.
Jason Dunham, Charlie Corrarino, and Desiree Tullos, my committee members,
provided endless expertise and brainstorming ideas for conducting research in
inhospitable locations; thank you for all of the feedback and intellectual support on
these projects.
I simply don’t have enough room to properly thank all the individuals and
organizations that helped me with the complex tasks of designing and completing two
separate fieldwork-intensive projects; so many people helped with the planning,
funding, permitting, field work, data collection, lab work, data analysis, historical
research, and editing that listing each name would fill pages upon pages. However,
there are some people who simply cannot remain in the shadows. Eric Baxter, Ed
Stephan, Jena Christiansen, and Gabe Dour were instrumental in the planning and
execution of both projects. I also thank Rich Grost, Michael Swanson, Bert Ortiz, Pat
McCrae, Dean Finnerty, Eric Figura, Bryan Deck, Gary Rowe, countless
Steamboaters, flyfishers, and the OSU Fisheries & Wildlife Club for all the
brainstorming and special efforts to capture wild steelhead on the North Umpqua
River; those months spent chasing elusive steelhead were not in vain. I appreciate the
friendships that sprouted along the river’s edge. Holly Huchko, Laura Jackson, Dave
Harris, Sam Moyers, and Dan Meyer of ODFW-Roseburg were indispensable in the
planning and permitting of the telemetry work. I give thanks for Ralph Lampman, who
helped me with both telemetry setup and trouble shooting. The local, historical
knowledge gained from Jeff Dose, Frank Moore, Craig Street, and countless others is
priceless; there are many things you cannot learn from a book.
Projects such as these cannot be accomplished without substantial financial
support and investments in both time and equipment; I thank the staff of the Umpqua
National Forest, Oregon Dept of Fish & Wildlife, Oregon Hatchery Research Center,
PacifiCorp, and the Federal Highways Administration for the financial support,
collaboration, and cooperation over the duration of the project. Scholarships from
Oregon State University, the North Umpqua Foundation, the Federation of Fly
Fishers, and several others helped pay for tuition, lodging, and fuel to travel back and
forth weekly between campus and Toketee for several years. Telemetry and fish
transport equipment was graciously loaned by ODFW, specifically the Oregon
Hatchery Research Center, Rock Creek Hatchery, Alsea Hatchery, and the Oregon
State University Department of Fisheries & Wildlife.
I wish to thank my family and close friends who helped me wade through grad
school, field projects, overgrown forests, dirty creeks, and other dark places. Finally, I
wish to thank my husband, Ray Lewis, for keeping me warm, fuzzy, and sane through
this whole process; my life would not be the same without you. Thank you!
“I sit in happy meditation on my rock, pondering, while my line dries
again, upon the ways of trout and men. How like fish we are: ready,
nay eager, to seize upon whatever new thing some wind of
circumstance shakes down upon the river of time! And how we rue our
haste, finding the gilded morsel to contain a hook.”
–From A Sand County Almanac and Sketches Here and There by Aldo
Leopold
CONTRIBUTION OF AUTHORS
David L.G. Noakes (Oregon Hatchery Research Center / Department of
Fisheries and Wildlife / Oregon State University) contributed to the study design, data
analysis, and editing of all sections of this manuscript. Jennifer McKay (CEOAS
Stable Isotope Lab / Oregon State University) contributed to the lab analysis of all the
isotopic analyses and revision of both manuscripts. Bruce Morrison (Ontario Ministry
of Natural Resources, Lake Ontario Fisheries Management Unit, Glenora Fisheries
Station) provided modeling expertise for the mixed-effects modeling work on stable
isotopes in this manuscript. Eric Baxter, Jena Christiansen, Gabe Dour, and Ed
Stephan assisted with all aspects of sample and data collection for all sections of this
manuscript.
TABLE OF CONTENTS
Page
CHAPTER 1: INTRODUCTION .................................................................................. 1
STUDY OBJECTIVES .......................................................................................... 3
GENERAL METHODS ......................................................................................... 5
STUDY LOCATION ............................................................................................. 7
CHAPTER 2. BEHAVIOR, HOLDING, AND OUTMIGRATION PATTERNS
OBSERVED IN WILD SUMMER STEELHEAD, O. MYKISS, ABOVE A
HYDROPOWER DAM IN THE HEADWATERS OF THE NORTH
UMPQUA RIVER ........................................................................................... 12
ABSTRACT ......................................................................................................... 13
BACKGROUND.................................................................................................. 14
METHODS .......................................................................................................... 18
RESULTS AND DISCUSSION .......................................................................... 20
CONCLUSIONS AND RECOMMENDATIONS .............................................. 24
REFERENCES ..................................................................................................... 28
CHAPTER 3: RESTORING ANADROMY: IDENTIFYING HISTORICAL FISH
PASSAGE BEYOND NATURAL OBSTACLES IN THE OREGON
CASCADES WITH STABLE ISOTOPE ANALYSIS ................................... 32
ABSTRACT ......................................................................................................... 33
INTRODUCTION................................................................................................ 33
MATERIALS AND METHODS ......................................................................... 35
RESULTS ............................................................................................................ 45
DISCUSSION ...................................................................................................... 55
REFERENCES ..................................................................................................... 60
TABLE OF CONTENTS (CONTINUED)
Page
CHAPTER 4: GENERAL DISCUSSION ................................................................... 64
CHAPTER 5: BIBLIOGRAPHY................................................................................. 67
CHAPTER 6: APPENDICES ...................................................................................... 76
APPENDIX A: TELEMETRY RESULTS .......................................................... 77
APPENDIX B: STATISTICAL ANALYSES ..................................................... 85
APPENDIX C: MIXED EFFECTS MODELING ............................................... 90
APPENDIX D: SUBSET TRENDS..................................................................... 96
APPENDIX E: STABLE ISOTOPE PHOTOS ................................................. 103
LIST OF FIGURES
Figure
Page
2.1. Map of North Umpqua River telemetry study area .............................................. 16
2.2. Location of O. mykiss tag detections in the North Umpqua River ...................... 22
3.1. Foliar transect locations in the North Umpqua River Basin ................................ 38
3.2. Foliar δ15N depositional trends in fitted mixed-effects model ............................. 47
3.3. Final fitted model showing Foliar δ15N intercepts ............................................... 48
3.4. Final fitted model showing Foliar δ15N response to historical salmon presence . 49
LIST OF TABLES
Table
Page
2.1. Summary of summer steelhead tracking from 6/23/2010 to 5/19/2011 ............... 23
3.1. Distances between upper and lower foliar nitrogen transects .............................. 40
3.2. Characteristics and mean foliar δ15N of transects ................................................ 42
3.3. Comparison in absolute foliar δ15N values between bankside and upslope trees . 50
3.4. Frequency counts of δ15N depositional trends in SW Oregon ............................. 53
3.5. C:N ratios observed in the North Umpqua River basin ....................................... 54
RECONNECTING AQUATIC HABITATS: VALIDATING HISTORICAL
HABITAT USE BY ANADROMOUS FISHES USING TELEMETRY AND
STABLE ISOTOPES ABOVE BARRIERS
CHAPTER 1: INTRODUCTION
Deciding to provide fish passage around barriers, or to completely remove
hydropower dams to restore habitat access for migratory fishes is a contentious, and
often litigious, global issue. As migratory fish populations have continued their
general downward trends, a great deal of research has been conducted to understand
the biological impacts of man-made obstacles and barriers. Almost every discussion is
focused on the measurable, ecological benefits to be gained versus the economic costs
to both stakeholders and the general public. Restoring aquatic habitat connectivity for
migratory fishes in coastal rivers can be an expensive approach to restoration if the
total length of river to be reconnected is uncertain because of natural or anthropogenic
barriers.
Obstacles that hinder fish movement may be dependent upon flow, whereas
barriers prevent fish movement under all flow conditions. Natural physical features
such as bedrock falls, chutes, cascades, or extreme velocities can also prevent fish
migration (Castro-Santos 2006, Pess et al. 2012, Pess et al. 2011, Powers and Orsborn
1985). Obstructions may be partially passable by certain individuals or temporarily
passable at specific times of the year under certain hydrological or temperature
conditions (Larinier 2000).
2
Effective mitigation projects require documentation of fish successfully
passing obstacles and re-colonizing substantial restored habitat. Man-made barriers
can be removed or modified to allow fish passage. These are often very expensive
endeavors, and the benefit is often limited by the existence of additional obstacles or
barriers further upstream (Anderson and Quinn 2007, Bartholomew et al. 2007,
Brenkman et al. 2012, Burke et al. 2008, Moser et al. 2011, Robards and Quinn 2002,
Roni et al. 2002). It has been common to modify natural features to facilitate fish
passage, but that practice is less common now due to growing respect for natural
partitions of aquatic communities. This balance between the cost of modifying manmade barriers and the inability or unwillingness to modify natural ones emphasizes a
need to better understand fish passage at natural features and to better estimate the
benefits of providing passage at man-made ones (Katopodis and Williams 2012).
Radio telemetry has often been the standard for determining both habitat usage
and fish passage (Sisak and Lotimer 1998). However, telemetry is both time and labor
intensive and results are generally site-specific. Capturing fish and releasing them
above potential barriers is considered an acceptable measure during the first years of
active restoration programs (Larinier 2000), and telemetry is the technique of choice to
validate those programs. A second, complimentary technique to address questions of
barriers and fish passage is the analysis of stable isotopes within ecosystems. Stable
isotope analysis can be an effective method to determine historical nutrient linkages
between terrestrial and aquatic ecosystems, but deposition patterns in riparian habitats
can be affected by biological and anthropogenic processes (Bilby et al. 2003, Bilby et
3
al. 1996, Drake and Naiman 2007, Drake et al. 2011, Helfield and Naiman 2003,
Hocking and Reimchen 2009, Hocking and Reynolds 2011, Holtgrieve and Schindler
2010). Each method has its challenges, and resource managers are often left
wondering which technique would provide the best information specific to their
watershed. My research combined both telemetry and stable isotope analyses to reduce
the uncertainties surrounding fish behavior and fish passage in restored habitat above a
functioning hydropower dam in the headwaters of the North Umpqua River, Oregon.
STUDY OBJECTIVES
The main goal of this project was to provide management recommendations
for habitat restoration projects upstream of the Soda Springs Dam and above a natural
waterfall in the Fish Creek tributary on the North Umpqua River in Oregon. I tested
the hypothesis that the historical distribution of anadromous salmonids would have
extended upstream from Soda Springs Dam to the base of Toketee Falls in the mainstem North Umpqua River and throughout the Fish Creek sub-basin. To test the
marine-nitrogen subsidy modeling hypotheses, I used a linear mixed-effects model
(Koyama et al. 2005) that had previously identified both salmon-bearing and nonsalmon bearing streams in Idaho.
I used radio telemetry to gather spatial distribution data on relocated,
anadromous summer steelhead, O. mykiss, as they moved into the restored habitat,
spawned, and outmigrated from the study area above Soda Springs Dam. I collected
information concerning the timing of holding, spawning, and kelt outmigrations to
guide resource management efforts to successfully pass fish both upstream and
4
downstream of the hydropower dam via a newly-constructed fish ladder. I predicted
that tagged steelhead would move throughout the entire restored area above Soda
Springs Dam and that some of the tagged steelhead would move upstream into the
Fish Creek tributary and attempt to pass the 4.2 m high natural waterfall obstacle 5.1
km upstream. I also predicted that there would be no delays in the timing of spawning
activities in tagged fish above Soda Springs Dam relative to fish spawning naturally
below Soda Springs Dam.
I used stable isotope analyses to detect marine-nutrient inputs as a way of
determining what habitats fish utilized prior to the building of Soda Springs Dam. I
collected Douglas fir foliage and riparian soils on transects located in both historically
anadromous and non-anadromous streams to document nitrogen deposition trends
above and below both anthropogenic and natural obstacles. I predicted that there
would be significant differences in foliar δ15N enrichment and deposition patterns
above and below waterfalls similar to the results observed in the Snake River basin in
central Idaho (Koyama et al. 2005). I predicted that I would observe enriched δ15N
foliar values in historical salmon-bearing streams when compared to depleted δ15N
foliar values in historical non-salmon-bearing streams in the riparian habitats of the
North Umpqua River. I predicted that δ15N foliar values would decrease from
bankside to upslope locations in salmon-bearing streams. I also predicted that there
would be no detectable gradient in δ15N foliar values for those streams historically
devoid of salmon. Finally, I predicted that I would be able to identify the habitat
5
upstream of the 4.2 m high waterfall obstacle in Fish Creek as either historically
salmon-bearing or non-salmon bearing based upon the presence or absence of an
enriched gradient for δ15N foliar values.
GENERAL METHODS
Telemetry
All procedures for this study were conducted with the approval of the
Institutional Animal Care and Utilization Committee (IACUC) of Oregon State
University (ACUP # 4045) and the National Oceanic and Atmospheric Administration
(NOAA) Fisheries and National Marine Fisheries Service (Federal 4-d Permits #
15290 and 15482). The Oregon Department of Fish and Wildlife approved fish
transport to the release site upstream of Soda Springs Dam. I collected fish using one
of two main methods, hook and line with artificial lures in the Wild and Scenic River
section of the North Umpqua River, and fish salvages conducted by PacifiCorp and
other agencies immediately below Soda Springs Dam prior to in-stream construction
work for the fish ladder. I recorded water temperature, date, handling time, species,
sex, condition, length, and any distinguishing marks (fin damage, scars, net, or claw
marks, etc.) for each fish that I handled. Fish judged to be in good physical condition
were kept for tagging as representative of the age class of interest (based on size).
Hatchery steelhead were counted and released downstream without further processing.
All other fish species were counted and released without any further processing
beyond species identification.
6
I conducted all animal procedures on the stream bank downstream of Soda
Springs Dam prior to loading for transport. I brought fish to shore in a landing net and
then placed them in aerated river water in a live well for further processing. I
identified each fish for species and apparent sex before photographing it (Olympus
850SW digital camera) to show overall body condition and distinguishing
morphological characteristics. I released fish back into the North Umpqua River above
Soda Springs Dam and the Soda Springs Reservoir directly across from the Slide
Creek Powerhouse after being photographed, measured for fork length on a measuring
board (nearest mm), radio-tagged, and double Floy tagged to continue their upstream
migration. I used LOTEK SR-M16-25 tags (16mm x 50mm in size with a 30cm long
external antenna) in this study. Each tag weighed 17 grams in air and had a variable
burst rate of 4.5 to 5.0 seconds and a motion sensor. I tagged each fish with one
orange and one white Floy tag. All tags displayed contact information for the Umpqua
National Forest. I collected non-lethal genetic tissue samples with a 0.5 mm punch to
the dorsal lobe of the caudal fin. Samples were stored in vials (95% ethanol) or in
paper envelopes as dry samples (Schreck and Moyle 1990). All tissue samples are
stored at the Oregon Hatchery Research Center, Alsea, Oregon.
Stable Isotopes
I used stable isotope analysis of conifer foliage in both historically anadromous
and non-anadromous streams and rivers (Koyama et al. 2005). Positive (anadromous)
controls were defined as areas with current migratory salmonid populations. Negative
(non-anadromous) controls were defined as areas upstream of well-documented,
7
impassable natural waterfalls. Unknown areas were all located above the 4.2 m high
natural waterfall 5.1 km upstream in Fish Creek. All transects were located in mid- to
late-successional stage forests.
STUDY LOCATION
The confluence of the North Umpqua River and the main-stem Umpqua River
begins at 179 River Kilometers (RKm) inland on the western slopes of the central
Cascade Mountains in southern Oregon. The North Umpqua River drains 3,520 km2
(Wallick 2011) and is legendary for its recreational fishing opportunities, rugged
basalt river canyons, and natural beauty. Winchester Dam is located 11.2 Rkm
upstream on the North Umpqua River just north of Roseburg, Oregon and is a known
obstacle for some fish species. More than 300 (Rkm) inland on the North Umpqua
River, a series of dams known collectively as PacifiCorp Energy’s (PacifiCorp) North
Umpqua Hydroelectric Project (NUHP) were built in the 1950s. NUHP occupies about
1,400 hectares on the Umpqua National Forest and consists of eight dams, eight
powerhouses, several reservoirs, and about 70 km of waterways. Most of the NUHP is
located upstream from the 40 meter (m) high Toketee Falls that is the historic barrier
to anadromous fish migration on the North Umpqua River at approximately RKm 120.
However, the 25 m high Soda Springs Dam at RKm 112, downstream of Toketee
Falls, is the second highest dam in the project and has blocked all upstream migration
of fish since its construction in 1952 (PacifiCorp 1995). Between Toketee Falls and
Soda Springs Dam along the main-stem North Umpqua River is Fish Creek at Rkm
115.5. Fish Creek is a substantial tributary to the North Umpqua River, and it has a 4.2
8
m high waterfall complex 5.1 km upstream that may be either an obstacle or a barrier
to fish passage. This 4.2 m high waterfall was the focus for my study.
On October 28, 1988, a 54 km section of the river immediately downstream of
the Soda Springs powerhouse down to Rock Creek was designated as a Wild and
Scenic River (WSR) as part of the National Wild and Scenic Rivers System created by
Congress in 1968 (Public Law 90-542; 16 U.S.C. 1271) (USDA Forest Service and
USDI Bureau of Land Management 2002). As part of the Settlement Agreement for
the NUHP Federal Energy Regulatory Commission (FERC) relicensing in 2003,
PacifiCorp completed a fish ladder over Soda Springs Dam in November 2012 to
reconnect the WSR section of the North Umpqua River and re-establish access for
adult anadromous fish to move above Soda Springs Dam up to the Slide Creek Dam
(RKm 117.8) and within the lower section of Fish Creek at least to the base of the 5.1
km waterfall complex. An estimated 40 km of aquatic habitat will be accessible to
those species capable of successfully passing the Soda Springs fish ladder if fish can
pass the 5.1 km natural obstacle in Fish Creek (USDA Forest Service 1997). If fish
cannot pass the obstacle at 5.1 km, then the reconnected aquatic habitat is limited to
about 10.5 km (PacifiCorp 1998).
Several anadromous species, including spring and fall Chinook (Oncorhynchus
tshawytscha), coho (O. kisutch), coastal cutthroat trout (O. clarkii), summer and
winter steelhead (O. mykiss), and Pacific lamprey (Entosphenus tridentatus) occur in
the North Umpqua River year round at various stages in their life cycles. All
anadromous fish species in the Umpqua River Basin have experienced substantial
9
declines in historical abundance due to hydropower development and operations,
commercial and recreational harvest, hatchery supplementation, and habitat alteration
since European settlement in the mid-1800s (Coates 2012). Historical estimates based
on cannery data, Native American harvest records, and anecdotal evidence vary by an
order of magnitude for the species still found in the river basin (Lichatowich 1999,
Meengs and Lackey 2005). Fish count data from Winchester Dam are helpful for
documenting adult returns since 1945 (Huchko 2012), but most fish populations had
already been declared as overharvested by the early 1900s (Lichatowich 1999, Meengs
and Lackey 2005). For example, Meengs and Lackey (2005) estimated Umpqua
historical returns for coho of almost 200,000 individuals and 20,000 Chinook based
upon cannery record extrapolation. Dam counts from 1945 to 2012 indicate
approximate mean returns of 10,000 spring Chinook, 500 fall Chinook, 4,100 coho,
7,000 summer steelhead, 5-10,000 winter steelhead, 50 Pacific lamprey (reduced from
a peak of approx. 45,000 in the 1960s) (Lampman 2011), and fewer than 100 sea-run
cutthroat annually (Huchko 2012).
The North Umpqua River Basin generally experiences mild, wet winters and
xeric summers. Within the Umpqua National Forest, the annual precipitation ranges
from 86 centimeters (cm rainfall equivalent) at the lowest elevations to 183 cm in the
extreme headwater areas on the southern end of the Forest boundary. The north end of
the Forest boundary receives between 127 and 203 cm of rainfall equivalent at the
highest elevations. The majority of the annual precipitation (80%) falls between
October 1 and March 31; July and August are typically the hottest and driest months.
10
Snow usually persists throughout the winter months above 900 m elevation (USDA
Forest Service 1976).
Within the study area (from Glide, OR up to the Rogue-Umpqua Divide near
Diamond Lake), all of the streams are in mountainous reaches with highly constrained
stream channels; none of the streams in the study area can be characterized as
meandering spawning reaches with finer textured soils. Approximately 20% of the
North Umpqua River Basin lies above 1,700 m elevation and is thus primarily snowmelt derived, delivering a larger percentage of groundwater for summer base flows
than does the South Umpqua River Basin at consistently lower elevations (PacifiCorp
1998). Rain-on-snow events do occur and occasionally create higher-than-average
flows in December and January. Soils are generally deep, porous pumice. Vegetation
includes Oregon white oak (Quercus garryana), tanoak (Lithocarpus densiflorus),
chinkapin (Castanopsis chrysophylla), western hemlock (Tsuga heterophylla),
Douglas fir (Pseudotsuga mensiesii), western red cedar (Thuja plicata), bigleaf maple
(Acer macrophyllum), and red alder (Alnus rubra). The entire study area is federal
forest land, managed for multiple purposes including recreation, forest products, and
timber. The Umpqua River is comparable to other Pacific Rim rivers with ecologically
intact floodplains, decreasing salmonid populations, human presence, and habitat
degradation (Coates 2012).
This thesis is presented in manuscript form. Chapter 2 investigates steelhead
holding, spawning, and outmigration behaviors in restored habitat above a functioning
hydropower dam over a one - year period. Chapter 3 investigates foliar δ15N
11
deposition patterns above and below migration barriers in the headwaters the North
Umpqua, of one of the longer coastal rivers in Oregon. Chapter 4 provides conclusions
and a general discussion of both the telemetry and stable isotope projects, and
management recommendations.
12
CHAPTER 2. BEHAVIOR, HOLDING, AND OUTMIGRATION PATTERNS
OBSERVED IN WILD SUMMER STEELHEAD, O. MYKISS, ABOVE A
HYDROPOWER DAM IN THE HEADWATERS OF THE NORTH UMPQUA
RIVER
Sierra K. Lewis
Department of Fisheries and Wildlife, Oregon State University, 104 Nash Hall,
Corvallis, Oregon 97331
In preparation for Animal Biotelemetry, BioMed Central, London, United Kingdom
13
ABSTRACT
I conducted a study to identify preferred spawning habitat for anadromous
summer steelhead (Oncorhynchus mykiss) above a 60-year-old hydropower dam in the
headwaters of the North Umpqua River, Oregon. Like many other historical salmonbearing rivers, little documentation exists for anadromous fish presence above
potential natural obstacles upstream of Soda Springs Dam. The prediction was that if
migratory salmonids are allowed to move upstream of the dam using a new fish ladder
they would utilize newly available upstream spawning habitat. I captured, radiotagged, and transplanted wild adult summer steelhead, O. mykiss, above Soda Springs
Dam; and I tracked daily movements for 1 year to observe habitat preferences. I
documented holding locations, the timing of spawning activities, and the outmigration
of kelts. Most of the tagged adults showed incremental daily movements near the
release site or downstream into the hydropower reservoir but did not travel further
than 1 kilometer upstream in the main river channel. There was no evidence from
telemetry that any fish moved upstream in Fish Creek, the newly accessible habitat of
interest in this project. I recorded movements and localized activities that suggested
some of the fish spawned and subsequently moved downstream towards the Pacific
Ocean, but I collected evidence that spawning behaviors of tagged fish above Soda
Springs Dam were delayed by 2 to 3 weeks, relative to fish spawning naturally below
the dam. My data suggest that steelhead will use restored habitat above dams, but that
re-colonization behaviors are variable and may be affected by altered flow regimes
within the restored habitat upstream of the dam amongst other variables.
14
BACKGROUND
Predicting salmonid habitat usage patterns above hydropower dams after
decades of complete absence is a difficult task. Installation of a fish ladder, or
breaching a dam to reconnect anadromous habitat creates new challenges for resource
managers when questions arise concerning historical passage above natural obstacles
upstream of the modified barrier (Burke et al. 2008, English et al. 2006, Keefer et al.
2008, Pess et al. 2012). Natural physical features such as bedrock falls, chutes,
cascades, or extreme velocities can prevent fish migration (Castro-Santos 2006, Pess
et al. 2012, Pess et al. 2011, Powers and Orsborn 1985), so most managers recommend
telemetry studies at natural obstacles to better estimate the benefits of providing
passage at man-made ones (Katopodis and Williams 2012). I conducted a 1-year
telemetry study in the headwaters of the North Umpqua River in southwest Oregon to
address this practical management question prior to the installation of a fish ladder.
Like many other salmon-bearing rivers in the Pacific Northwest with recently
restored habitat, little historical documentation exists for anadromous fish presence in
the uppermost reaches of North Umpqua River. Several species of native, migratory
fish, including spring Chinook (Oncorhynchus tshawytscha), coho (O. kisutch), coastal
cutthroat trout (O. clarkii), summer and winter steelhead (O. mykiss), and Pacific
lamprey (Entosphenus tridentatus), travel more than 300 kilometers (km) inland from
the Pacific Ocean before they reach the base of the 60 year old, 25 meter (m) high
Soda Springs Dam (SSD) on the North Umpqua River. As part of the 2003 relicensing agreement for the North Umpqua Hydropower Project (NUHP), which
15
includes SSD, a fish ladder was installed at SSD and made ready for operation as of
November 2012. Most of the NUHP is located upstream from the 40 meter (m) high
Toketee Falls that is the historic barrier to anadromous fish migration on the North
Umpqua River at approximately RKm 120 (Appendix E). However, the 25 m high
Soda Springs Dam at RKm 112, downstream of Toketee Falls, is the second highest
dam in the project and has blocked all upstream migration of fish since its construction
in 1952 (PacifiCorp 1995). Between Toketee Falls and Soda Springs Dam along the
main-stem North Umpqua River is Fish Creek at Rkm 115.5. Fish Creek is a
substantial tributary to the North Umpqua River, and it has a 4.2 m high waterfall
complex 5.1 km upstream that may be either an obstacle or a barrier to fish passage.
This 4.2 m high waterfall was the focus for my study (Figure 2.1).
Figure 2.1. Map of North Umpqua River telemetry study area showing stationary receivers (flags), capture sites (Xs), release
location (triangles), waterfalls (stars), and hydropower dams (zigzags). Solid stars indicate waterfalls impassable to any fish
species. Note the location of Toketee Falls in relation to the two main-stem dams downstream (Soda Springs Dam and Slide
Creek Dam).
16
17
Fish Creek is a substantial tributary to the North Umpqua River, and it has a
4.2 m high waterfall complex that may be either an obstacle or a barrier to fish passage
(5.1 km upstream) (Appendix E). An estimated 40 km of aquatic habitat is accessible
to species capable of successfully passing both the Soda Springs fish ladder and this
waterfall complex in Fish Creek (USDA Forest Service 1997). If fish cannot pass the
obstacle at 5.1 km, then the reconnected aquatic habitat is limited to about 10.5 km
(PacifiCorp 1995, 1998). Summer and winter steelhead (O. mykiss) are predicted to be
the species most likely to successfully navigate the 4.2 m high obstacle in Fish Creek
due to their natural leaping capabilities. Summer steelhead arrive at SSD in June and
typically remain until January of the following year (Street 2010). Spawning occurs
from January to February, after which kelts emigrate. Winter steelhead arrive in
February, spawn, and then kelts emigrate by May of the same year. Management
concerns included differential use of the available habitat by summer and winter
steelhead as well as varying hydrologic flows that may affect passage efforts at the
waterfall complex.
Study Objectives
The main objective of this project was to provide management
recommendations for habitat restoration projects upstream of the Soda Springs Dam
and above the waterfall complex. Additional objectives included (1) capture and
release of wild, iteroparous O. mykiss above Soda Springs Dam prior to completion of
the fish ladder (Larinier 2000), (2) stream-by-stream analysis of preferred habitat for
spawning, (3) documentation of holding, spawning, and outmigration timing for
18
steelhead above SSD, and (4) documentation of migration delays due to natural or
anthropogenic obstacles in Fish Creek. Ideally, fish would utilize the entire 40 km
stretch of newly available habitat. I also predicted that there would be no delays in the
timing of spawning activities in tagged fish above Soda Springs Dam relative to untagged fish spawning naturally below Soda Springs Dam.
METHODS
For this study area, I programmed three stationary LOTEK SRX600 receivers
to reject tag IDs not specifically deployed on a single radio frequency due to
substantial electronic interference. I maximized antenna detection efficiencies by
physically testing a deployed active radio tag (LOTEK SR-M16-25, 16mm x 50mm in
size with a 30cm long external antenna, 17 grams in air) mounted on a wading staff
held continuously under water. Each station had at least two 4 or 6-element Yagi
antennas arranged to detect upstream and downstream movements; the Fish Creek
receiver had a third antenna placed to detect movements in and out of the tributary
mouth. North Umpqua River flows from October 2010 to June 2011 were higher than
anticipated due to NUHP reduced holding capacities during fish ladder construction
and repeated, seasonal high water events, so I conducted redd surveys when possible
(U.S. Geological Survey 2013).
All fish and radio-tracking data were collected by tracking 14, wild summer
steelhead from June 2010 to late May 2011 after initial capture, tagging, and
relocation. Four fish were caught using hook and line over 531 hours between July and
November 2010; the other 10 fish were obtained from salvage operations immediately
19
below SSD (RKm 112) between June and December 2010 (Appendix A). I recorded
water temperature, date, handling time, apparent sex, condition, length, and any
distinguishing marks for each fish. I gastric-tagged, Floy tagged, measured fork length
(FL), photographed, and collected a caudal-punch genetic sample from each wild
steelhead (Table 2.1). I released 14 tagged O. mykiss upstream of the SSD reservoir
(RKm 114.5). During multiple fish salvages at Soda Springs Dam, I documented
hatchery stray abundances for spring Chinook (30%), coho (% not recorded), summer
steelhead (40%), and South Umpqua winter steelhead (1 kelt observed).
Because each radio tag had a unique ID and audio signal, I tracked fish in real
time. I gathered daily spatial distribution data as tagged fish moved within the mainstem North Umpqua, spawned, or left the study area above SSD. Tagged transplant
fish were assumed to behave as wild fish with no observed differences in behavior or
physical performance (Keefer et al. 2004, Lucas 1989, Lucas and Johnstone 1990,
Martinelli et al. 1998, Neely et al. 2009, Smith et al. 2009, Welch et al. 2007).
Data Analysis
I combined all telemetry data collected from the fixed and mobile receivers
into a single spreadsheet to archive the dates and locations of each tagged fish. I
examined the data, identified signals from individual fish, and removed false
detections. I documented fish as present in a given location only if the tag signal
strength was above 100 (out of possible 225 units), if an audible signal was observed
during mobile tracking, or if numerous tag detections occurred for a given ID over
several consecutive minutes at any of the stationary receivers.
20
RESULTS AND DISCUSSION
Movement Patterns – Exploration, Holding, Spawning, Emigration
Tagged fish did not show any extensive movement in the river or tributaries
above the dam and held in deep, fast flowing water ≥4m in depth near large
submerged features such as boulders, bedrock ledges, or overhanging riverbanks
(Appendix A). Most of the tagged adults showed incremental daily movements near
the release site or downstream into the hydropower reservoir but did not travel further
than 1 kilometer upstream in the main river channel from the release site (Appendix
A). Fish 17 outmigrated from the study area in January 2011 during a high flow event;
but radio tag movements ceased just upstream of Boulder Creek within a few days and
never continued downstream. Fish 19 died within hours of salvage and tagging
procedures; the radio tag was re-deployed in another fish which was subsequently
poached from Soda Springs Reservoir in late March 2011. At the conclusion of
tracking efforts in late May 2011, 5 tagged fish remained above Soda Springs Dam; all
tag IDs indicated fish were alive and moving upstream between SSD reservoir and the
Slide Creek powerhouse (Rkm 114.5).
I also recorded continuously mobile tag signals and highly localized activities
from 3/16/11 to 3/23/11 50m upstream of the Fish Creek confluence in the main-stem
North Umpqua River that suggested three fish spawned on a very small gravel patch
and subsequently moved downstream towards the Pacific Ocean; these same data
suggested that spawning behavior of tagged summer steelhead above SSD was
delayed by several weeks relative to untagged, summer steelhead spawning below the
21
dam. Three of the four confirmed outmigrating fish traveled approx. 100 RKm
downstream in 72 – 96 hours before signals were lost downstream of Winchester Dam
due to electronic interference and lack of access to the river for tracking (Figure 2.2).
Between October 2010 and March 2011, three significant peak river flows
were documented at US Geological Survey gauge station # 14315950 located 10 km
upstream in Fish Creek, including one 7-year flooding event. All three peak flows
increased from average seasonal flows of about 2.8 m3/second up to a maximum of
56.6 m3/second and subsided without any significant changes in behavior or location
from the tagged fish (Appendix A).
Figure 2.2. Location of O. mykiss tag detections in the North Umpqua River from January to May 2011 during outmigration.
Steelhead were individually tracked as they passed over Soda Springs Dam during unplanned spill events until radio signals
were lost near Winchester Dam 100 Rkm downstream.
22
Table 2.1. Summary of summer steelhead tracking from 6/23/2010 to 5/19/2011. a = Soda Springs Dam. *=Suspected
Emigration Date. Two tags malfunctioned, allowed for intermittent tracking, and could not be confirmed as present/absent at
the conclusion of the study.
23
24
CONCLUSIONS AND RECOMMENDATIONS
My data suggest that steelhead will use restored habitat above dams, but that
re-colonization behavior is variable and may be affected by altered flow regimes
within the restored habitat upstream of the dam or a variety of other site-specific
factors (Pess et al. 2012, Pess et al. 2011). Some observed behaviors, such as lack of
exploration, may be due to intra-specific density dependence, but they reflect current
habitat conditions and expected low population numbers above Soda Springs Dam
since I was able to capture and track approximately 14% of the total anadromous O.
mykiss population documented below the dam in average years (L. Jackson and S.
Moyers, ODFW 2009). Originally, I had planned on tagging 25 summer and 25 winter
steelhead but I could not obtain enough wild fish for tracking using the approved
methods (NOAA/NMFS 4-d Permits # 15290 and 15482).
Tagged summer steelhead preferred deep, fast flowing pools for holding
similar to other studies on North Umpqua summer steelhead (Baigún 2003). The three
tagged fish suspected of spawning moved to the much less prevalent, shallow (≤1m
deep), slower riffles with gravel substrate during suspected spawning activities. The
streambed of the North Umpqua River above Soda Springs Dam is typically
comprised of large boulders and bedrock ledges with very little gravel suitable for
building redds. The patch of suitably-sized spawning gravel 50m upstream of the Fish
Creek confluence occupied during suspected spawning activities was a direct result of
an un-related gravel augmentation and deposition study; no other gravel deposits were
observed during redd surveys in 2011 above the Fish Creek confluence. The majority
25
(>90%) of the gravel from the augmentation study was deposited downstream between
the Fish Creek confluence and Slide Creek Powerhouse (Street 2011) and that was
where the majority of tagged fish were documented for the duration of the study.
Anthropogenically-altered river flow regimes are of great concern if they
negatively affect habitat use and the timing of spawning behavior for declining
salmonid or migratory fish populations (Beeman et al. 2008, Bunt et al. 1999, Burke et
al. 2008, English et al. 2006, Keefer et al. 2008, Kurth 2013, Null et al. 2013). In Fish
Creek flows are diverted for hydropower once they pass a legally-determined
threshold (approximately 150 cfs). This results in water diversions typically occurring
during the higher flows from December until early summer each year. If fish passage
is flow dependent at the waterfall complex then chances of successful passage is
reduced by diversion of any amount of water when fish are present in the area. I
recommend that the historical environmental flow regime be restored to Fish Creek to
enhance re-colonization efforts.
Several species of native salmon, trout, and Pacific lamprey can now access
the restored habitat above Soda Springs Dam, I recommend additional telemetry
tracking be conducted on each species to identify habitats specific to each life history
stage (Anderson and Quinn 2007). If the overall timing for spawning is truly different
above Soda Springs Dam than below then new issues may be created regarding smolt
outmigration timing, predation, recruitment, and survival in the basin.
26
The continued presence and abundance of stray hatchery fish so far upstream
in the headwaters of this coastal Oregon river is cause for concern given the meager
wild adult returns (Araki et al. 2008, Araki and Schmid 2010, Berkeley et al. 2004,
Christie et al. 2011, Rand et al. 2012). I was not able to study hatchery steelhead
behavior in the restored habitat even though they will now have the same access
opportunities as wild returning adults. To prevent access to the restored habitat by
hatchery fish, I recommend installing a fish collection and sorting facility at the Soda
Springs Dam fish ladder (Fausch et al. 2009).
Current fishing regulations for the area are not adequate and were changed in
2013 to prevent all angler harvest of established, non-indigenous brown trout (Salmo
trutta) and brook trout (Salvelinus fontinalis) populations throughout the NUHP
(Oregon Department of Fish & Wildlife 2013). I recommend unlimited harvest of
brown and brook trout for all of the main-stem North Umpqua River above Soda
Springs Dam and NUHP reservoirs to reduce inter-specific competition and predation
opportunities upon smolts. I recommend that Fish Creek receive the same protections
and regulations as all of the other tributaries in the WSR section of the river; this
would remove all angling pressure for both resident and migratory forms of O. mykiss.
Overall, my telemetry research suggests that further research is needed to
understand all of the biological and ecological impacts of the NUHP on native aquatic
species above Soda Springs Dam. I would recommend proceeding with extreme
caution before modifying the natural obstacle 5.1 km upstream in Fish Creek. I
recommend continued protection of existing late-seral riparian habitats, replacing
27
large wood in-stream (previously removed from area streams during stream “cleaning”
events in the 1960s), and re-introducing American beaver (Castor canadensis) to the
entire Umpqua National Forest. Beaver were historically present in the area and are
ecosystem engineers with documented benefits to salmonid populations. By seeking to
further clarify the habitat requirements for native aquatic species above the 4.2 m high
natural waterfall in Fish Creek, it may be possible to reduce the negative impacts of
invasive species, altered hydrological flows, and hatchery fish below the waterfall
while also improving the overall integrity of the local ecosystem.
Acknowledgements
I gratefully acknowledge the substantial support of Dr. David Noakes, my
major professor. I formally acknowledge the numerous financial grants, equipment
loans, and field assistance provided by the US Forest Service, Oregon State
University, Oregon Department of Fish and Wildlife, the Oregon Hatchery Research
Center, PacifiCorp, and the North Umpqua Foundation. I also acknowledge the
substantial guidance provided by ODFW personnel in the 4-d permitting process and
fish transport efforts over multiple field seasons; Tim Walters, Laura Jackson, Holly
Huchko, Joy Vaughn, Shivonne Nesbit, Dan Meyer, and Marc Garst all directly
contributed to the success of the telemetry project. Finally, I am thankful for the
fishing and radio-tracking efforts made over several months by Ed Stephan, numerous
volunteers, the North Umpqua Foundation, and the Steamboaters organizations of the
North Umpqua River.
28
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Castro-Santos, T. 2006. Modeling the Effect of Varying Swim Speeds on Fish Passage
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Christie, M.R., Marine, M.L., and Blouin, M.S. 2011. Who are the missing parents?
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Comparison of Adult Steelhead Migrations in the Mid-Columbia Hydrosystem
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Invasion versus Isolation: Trade-Offs in Managing Native Salmonids with
Barriers to Upstream Movement. Conservation Biology 23(4): 859-870.
Katopodis, C., and Williams, J.G. 2012. The development of fish passage research in a
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Keefer, M.L., Peery, C.A., Ringe, R.R., and Bjornn, I.C. 2004. Regurgitation rates of
intragastric radio transmitters by adult Chinook salmon and steelhead during
upstream migration in the Columbia and Snake Rivers. North American
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Oncorhynchus mykiss. Environmental Biology of Fishes 96(2-3): 355-362.
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Null, R., Niemela, K., and Hamelberg, S. 2013. Post-spawn migrations of hatcheryorigin Oncorhynchus mykiss kelts in the Central Valley of California.
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T.P. 2011. The Influences of Body Size, Habitat Quality, and Competition on
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2009. Evaluation of a Gastric Radio Tag Insertion Technique for Anadromous
River Herring. North American Journal of Fisheries Management 29(2): 367377.
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National Forest - Diamond Lake Ranger District, Idleyld Park, OR.
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Idleyld Park, OR.
31
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of steelhead trout (O-mykiss) surgically implanted with dummy acoustic tags.
Hydrobiologia 582: 289-299.
32
CHAPTER 3: RESTORING ANADROMY: IDENTIFYING HISTORICAL FISH
PASSAGE BEYOND NATURAL OBSTACLES IN THE OREGON CASCADES
WITH STABLE ISOTOPE ANALYSIS
Sierra Koch Lewis1, Jennifer McKay2, Bruce Morrison3, and David Noakes1,4
1
Department of Fisheries and Wildlife, Oregon State University, 104 Nash Hall,
Corvallis, Oregon 97331
2
College of Earth, Ocean, and Atmospheric Sciences, Oregon State University, Stable
Isotope Laboratory, 104 CEOAS Administration Building, Corvallis, Oregon 973315503
3
Ontario Ministry of Natural Resources, Lake Ontario Fisheries Management Unit,
Glenora Fisheries Station, Lake Ontario Management Unit, 41 Hatchery Lane, RR 4,
Picton ON K0K2T0
4
Oregon Hatchery Research Center, 2418 East Fall Creek Road, Alsea, Oregon 97324
In preparation for Canadian Journal of Fisheries and Aquatic Sciences
Canadian Science Publishing (NRC Research Press), Ottawa, Canada
33
ABSTRACT
We investigated evidence of historical anadromy throughout the North
Umpqua River basin in Oregon. We tested the hypothesis that we could identify
undocumented, historical salmon spawning reaches above obstacles using foliar δ15N
deposition patterns in Douglas fir trees, P. menziesii, (>100 years old). Our stable
isotope data indicated that the foliar δ15N deposition patterns were confounded by
elevation. We documented potential “salmon-derived” false positives on the Umpqua
National Forest above impassable waterfalls. Overall we found that foliar δ15N
deposition patterns had high inter and intra-site variability. When grouped by salmon
history, our data indicated a statistically significant negative correlation of foliar δ15N
values with historical salmon presence. Our linear mixed-effects modeling suggests
that the presence of salmon is the most important indicator of foliar δ15N values, rather
than the proximity of a sampled tree to stream flow above or below migratory barriers.
We were not able to identify previously undocumented salmon spawning habitat. Our
results from the foliar technique suggest that the method may not be as generally
applicable as has been suggested in the literature.
INTRODUCTION
Historically, mitigation projects to restore fish passage at man-made barriers
have defined project success as observed fish presence beyond the obstacle. Restoring
aquatic connectivity for migratory fishes in the headwaters of coastal rivers above
dams can be an expensive restoration proposition if the total length of stream or river
34
to be regained is uncertain because of natural obstacles. To detect salmon-derived
nutrient linkages and spatially quantify previously undocumented, historical salmonid
habitat within the headwaters of the North Umpqua River basin in Oregon we used an
accepted foliar isotopic technique (Koyama et al. 2005) above and below known and
potential obstacles and barriers. Obstacles that hinder fish movement may be
dependent upon flow, whereas barriers prevent fish movement under all flow
conditions. Natural physical features such as bedrock falls, chutes, cascades, or
extreme velocities can also prevent fish migration (Castro-Santos 2006, Pess et al.
2012, Pess et al. 2011, Powers and Orsborn 1985). Obstructions may be partially
passable by certain individuals or temporarily passable at specific times of the year
under certain hydrological or temperature conditions (Larinier 2000).
Anadromous species such as salmon act as nutrient vectors that contribute a
variety of marine-derived nutrients including isotopically heavy nitrogen (15N) and
carbon (13C) through the decay of their carcasses after spawning (Cederholm et al.
1999). In regions with substantial total biomass deposition due to salmon spawning
activities, these inputs of heavier isotopes can be detected using several methods at
different trophic levels (Bartz and Naiman 2005, Bilby et al. 2003, Bilby et al. 1996,
Finney et al. 2000, Hocking and Reimchen 2009, Janetski et al. 2009, Reichert et al.
2008). Fractionation of terrestrial nitrogen can lead to enriched 15N signals that are not
necessarily due to anadromous fish deposition (Perakis et al. 2006, Scott et al. 2009,
Scott et al. 2008). Historical populations and spatial distribution have also been
35
analyzed using marine derived nutrients detected in the riparian zone (Bilby et al.
2001, Drake and Naiman 2007, Garman et al. 2009, Gende et al. 2007, Gresh et al.
2000, Helfield and Naiman 2001, Helfield and Naiman 2002, Nagasaka et al. 2006).
Knowing historical distribution patterns is important for agencies seeking to
restore access to habitat used by anadromous fishes (Fellman et al. 2008, Fellman et
al. 2009, Janetski et al. 2009). Given the time scale of interest (60+ years of absence)
and low N-turnover rates, we selected long-lived Douglas fir for Nitrogen isotopic
analyses within the Umpqua National Forest. Following Koyama et al (2005), we
predicted that (1) the foliar δ15N values would be elevated bankside and would
decrease going upslope perpendicular to stream flow at sites with salmon present, (2)
there would be no elevated foliar δ15N values observed bankside at sites not accessible
to salmon, and (3) observed δ15N patterns at sites with unknown salmonid spawning
histories could be classified based upon existence or lack of elevated bankside δ15N
foliar values.
MATERIALS AND METHODS
Study Location
The confluence of the North Umpqua River and the main-stem Umpqua River
is at 179 River Kilometers (RKm) inland on the western slopes of the central Cascade
Mountains in southern Oregon (Figure 3.1); the North Umpqua River drains 3,520
km2 (Wallick 2011). At RKm 112, more than 300 Rkm inland, the 25 meter (m) high
Soda Springs Dam (SSD) has blocked a portion of historical salmonid habitat since
36
1952 (PacifiCorp 1995). SSD is the second highest dam in the series of 8 known
collectively as the North Umpqua Hydroelectric Project (NUHP). Most of the NUHP
is located upstream from the 37 m high Toketee Falls (Rkm 120) that is the known
historic barrier to anadromous fish migration on the North Umpqua River. Between
Soda Springs Dam and Toketee Falls along the main-stem North Umpqua River is
Fish Creek at Rkm 115.5. Fish Creek is a substantial tributary to the North Umpqua
River, and it has a 4.2 m high waterfall complex 5.1 km upstream that may be either
an obstacle or a barrier to fish passage (PacifiCorp 1998, USDA Forest Service 1997).
Determining whether or not fish passed upstream of this 4.2 m high waterfall was the
focus for our study. An estimated 40 km of aquatic habitat will be accessible to those
species capable of successfully passing the newly-constructed SSD fish ladder and the
waterfall complex at 5.1 km in Fish Creek (USDA Forest Service 1997). If fish
cannot pass the obstacle at 5.1 km, then the reconnected aquatic habitat is limited to
about 10.5 km (PacifiCorp 1998).
A priori sample collection
We collected 86 foliar samples from Douglas fir trees, P. menziesii, (>100
years old, diameter at breast height (DBH) = 1.25m ± 0.28) along 29 transects in the
North Umpqua River basin in mid- to late-successional forested areas within the
Umpqua National Forest in Oregon. We sited transects after careful interpretation of
topographic maps, GIS layers, ortho-corrected aerial photography, physical field
inspection, on-site stream gradient inspection, and visual confirmation of spawning
37
adults where possible to increase the likelihood of historical spawning events in these
tightly constrained, mountainous stream reaches (Figure 3.1). We sampled areas with
less than 5% of the understory foliar biomass from nitrogen-fixing plants (visual
estimates) when possible (Kohl and Shearer 1980).
We identified 13 of 29 anadromous (positive control) transects located
downstream of Soda Springs Dam with extant salmonid populations. We also included
the lower segment (first 5.1 km) of Fish Creek and a transect downstream of Toketee
Falls but above SSD. We identified 6 non-anadromous (negative control) transects
located upstream of Toketee Falls. Upper Boulder Creek, with a 15m high waterfall
downstream of Onion Creek confluence, was also defined as a negative control. We
selected 10 transects with undocumented salmonid history (unknowns) located
upstream of a 4.2 m high complex in the Fish Creek sub-basin. Distinctions between
barrier types were documented (e.g., large boulders, bedrock falls, debris jams, etc.).
We also calculated average soil Cation Exchange Capacity (CEC) values specific to
our study transects when available (USDA Forest Service 1976).
Figure 3.1. Foliar transect locations in the North Umpqua River Basin collected in 2011. White triangles are positive controls,
black triangles are negative controls, triangles with dots represent unknown historical salmonid transects, zigzags are dams,
and stars (white or black) represent documented waterfalls.
38
39
To address predictions 1 and 2, U.S. Forest Service certified tree climbers
collected the most-vigorous, current year foliage from the upper canopy (upper 1030% of total tree height) in full sun from bankside (0-10m), middle (10-50m), and
upper (100-150m) upslope. Ground crews recorded transect botanical species
composition, percent (of total site biomass) of N-fixing species, elevation, slope,
aspect, and collected soil samples adjacent to the bankside tree. Percent total biomass
of N-fixers observed within each transect was intentionally overestimated and
categorized as <1, <5, 5-10, and 15 percent to reduce potential signal masking by
terrestrial N sources. To account for transport of marine 15N by larger mobile
predators (Hilderbrand et al. 1999, Quinn et al. 2009), transects were sited 1-6
kilometers (km) above and below obstacles (Table 3.1). Transects above and below
obstacles had similar stream morphology, species composition, and site
characteristics. Transects were less than 30 m in width. Samples were kept cool during
transport and then frozen (-20oC) until isotopic processing.
Table 3.1. Distances between upper and lower foliar nitrogen transects (estimated via thalweg measurements in ArcMap v10).
Distance calculated between below Toketee Falls and the Clearwater River Falls is cumulative; for foliar δ15N signal to be
salmon-related, each fish would need to sequentially pass Toketee Falls, Whitehorse Falls, and Clearwater River Falls before
arriving at the foliar transect (total distance = 29.68 km). Some waterfalls in our study area, exceeding 4.5m in height, do not
block salmon passage.
40
41
Lab analyses
We analyzed 172 foliar subsamples (2 per tree) for δ15N, percent carbon (%C),
and percent nitrogen (%N). All nitrogen isotope data are expressed as δ15N values
(‰); air is the relative zero standard at 0‰ for δ15N. . Samples were dried to a
constant weight at 50oC for 48 hours, pulverized and homogenized using a Wigg-LBug machine, weighed and wrapped individually in tin cups (10.791 mg + 0.123 mg),
and then flash combusted at >1000oC using a Carlo Erba NA1500 Elemental
Analyzer. Following combustion CO2 was trapped and N2 was analyzed by
continuous-flow mass spectrometry on a DeltaPlus isotope ratio mass spectrometer at
the CEOAS Stable Isotope Lab at Oregon State University in Corvallis, Oregon.
International standards USGS-40 (glutamic acid, expected δ15N = -4.52‰), IAEA-N1
(ammonium sulfate, expected δ15N = +0.4‰), and leucine (observed δ15N= -0.26‰)
were run simultaneously for comparison. Mean δ15N values for all 172 foliar
subsamples ranged from -0.59 to -4.54 ‰ (Table 3.2).
Table 3.2. Characteristics and mean foliar δ15N of transects located in SW Oregon.
42
Table 3.2. (Continued) Characteristics and mean foliar δ15N of transects located in SW Oregon.
43
44
Post-hoc statistical analyses, including Linear-Mixed Effects Modeling
To address prediction 3 regarding classification of historical salmonid habitat
based upon existence or lack of elevated bankside δ15N foliar values, we assessed
several linear regression models with graphical diagnostics, metrics of data normality,
and residual plots (Appendix B) including variables for tree diameter at breast height
(DBH), salmonid history, elevation, aspect, gradient, percent N-fixers observed, foliar
and soil C:N ratios, human disturbance, and potential interaction between these terms.
We identified one outlier consistent across most of the variables analyzed. However
we did not remove this datum (Lower Copeland Creek – upslope tree) given that the
study’s objective was to compare positive and negative controls with unknown
transects to discern patterns. Removal of that value would not change the observed
bankside means, nor reduce the inter- and intra-site variability observed between
bankside, middle, and upslope trees. All data collected was transformed using natural
logarithms to improve normality and homoscedasticity. Because all foliar δ15N values
were negative, we transformed them by adding a constant value of 5 to each observed
data point prior to natural logarithmic transformation.
We modified Koyama’s (2005) original linear mixed-effects model since we
restricted sampling to a single conifer species (Douglas fir). This eliminated the
species variable and interaction term for DBH*Conifer species; we then added a new
variable for percent N-fixers (observed) and an interaction term for DBH*Salmon. To
balance the data set for linear mixed-effects modeling, the Lower Fish Creek, Lower
Steamboat – ALT, and North Umpqua River below Soda Springs Dam – North
45
transects were all removed because of missing data points. Using R statistical software
(version 2.15) and the lmer4 and lmerconvenience packages (R Development Core
Team 2012), we identified transect as a random effect and we used location as both a
fixed effect and also nested within transect as a random effect. We compared all
mixed-effect models tested in a backwards, stepwise approach using Akaike
Information Criteria (AIC) and p-values for each variable in the full model that
hypothesized: δ15Nitrogen ~ Salmon + DBH + Location + N-fixers + Salmon*DBH +
Salmon*Location + DBH*Location + Transect (random) + Location (nested within
Transect, random). We selected the final model based upon AIC values and backfitted
the most parsimonious model using REML (Appendix C).
RESULTS
Overall, we found that foliar δ15N deposition patterns were highly variable
(Figure 3.2), but with an unexpected, statistically significant negative correlation of
foliar δ15N values with historical salmon presence (Figures 3.3 and 3.4). Our linear
mixed-effects modeling suggests that the presence of salmon is the most important
indicator of foliar δ15N values, rather than the proximity of a sampled tree to stream
flow above or below migratory barriers (AIC= 116.4, F=6.3124, upper df=67, upper pvalue = 0.0031, lower df=37, lower p-value=0.0044, expl.dev (%)= 14.9019). We
were not able to identify any areas as salmon habitat using only foliar δ15N values. We
documented potential “salmon-derived” false positives on the Umpqua National Forest
46
above impassable waterfalls and observed no patterns in the absolute differences of
mean foliar δ15N values between bankside and upslope trees (Lower Watson and
Mowich Creek sites, Table 3.3).
47
Figure 3.2. Foliar δ15N depositional trends in fitted mixed-effects model (Foliar δ15N
~ Salmon only). A) Transects with no salmon historically. B) Transects where
salmon are still present currently. C) Transects with unknown salmonid history.
48
Figure 3.3. Final fitted model showing foliar δ15N intercepts in response to salmon
presence. Location is included in the figure but was not significant in the fitted model.
Without additional sampling, it cannot be conclusively stated that the unknown
transects are statistically similar to either the positive or negative control groups.
Figure 3.4. Final fitted model showing Foliar δ15N response to historical salmon presence. Vertical axis is the mean Foliar
δ15N natural logarithm; horizontal axis is documented salmon presence. Extrapolation or inferences to habitat with unknown
salmonid history is not possible using this technique until we can resolve the relationship between foliar δ15N deposition
patterns and current salmon populations.
49
50
Table 3.3. Comparison in absolute foliar δ15N values between bankside and upslope
trees for all transects sampled.
51
We observed several interesting, statistically significant trends within most of
the linear regression empirical models. However, comprehensive analysis using linear
mixed-effects modeling eliminated those explanatory variables and trends from the
final model. Detailed analyses using subsets of the foliar data within the linear mixedeffects model were not possible due to limited sample size. Specifically, empirical
subset analyses involving groups separated by observed depositional trends (Table
3.4), abundance / type of N-fixing species effects, and post hoc GIS human
disturbance data (Menke and Gamble 2012) (Appendix D) were not possible without
collecting additional foliar samples in the basin.
Soil Fertility and C:N ratios
Cation Exchange Capacity (CEC) values measured within the Umpqua
National Forest in 1976 (USDA Forest Service 1976), and sampled soils had low
fertility and low nutrient retention capabilities. We analyzed the original data and
calculated mean CEC and pH values for a subset of soil tests that were conducted
within the same sub-basins as our transect sampling. Anadromous sites had a mean
CEC = 25.15 and a pH=5.96 (n=16). Non-anadromous sites had a mean CEC =23.61
and a pH=6.25(n=10). Fish Creek sub-basin sites (Unknowns for this study) had a
mean CEC=15.59 and a pH=6.0(n=3). In another regional study, organic horizon CEC
values were reported at 76.2 (pH for “A” horizon samples = 6.52) for Drake and
Naiman’s salmon spawning sites (Drake and Naiman 2007). Organic horizon CEC
52
values were 11.6 (pH for “A” horizon samples = 5.89) for their non-salmon reference
sites. No pattern in CEC values was identified or attributable to salmon presence or
absence at the sites analyzed by Drake and Naiman.
C:N ratios calculated for our foliar and soil samples were highly variable and
sub-basin dependent (Table 3.5). C:N ratios for soil and foliar samples were not
confounded with elevational effects and allowed for interesting comparisons between
sub-basins, assumed salmon histories, and between trophic levels. For the soil samples
taken adjacent to the bankside foliar sample, the Unknown group mean ratio of 46.81
was significantly different than the positive or negative controls (p=0.004). The
Salmon Absent (negative controls) had a mean ratio of 26.98, and the Salmon Present
(positive controls) had a mean ratio of 28.55. The ratios of foliar samples for the
bankside trees were much more consistent. The Unknowns mean ratio was 48.35, and
the Salmon Negative and Positive control groups had a mean ratio of 42.52 and 40.54,
respectively.
Table 3.4. Frequency counts of δ15N depositional trends in SW Oregon for foliar δ15N transects. Unknown transects were
categorized as matching (decreasing δ15N upslope), not matching (increasing δ15N upslope, or exhibiting no trend (flat). Three
transects were not included because they lacked 3 datapoints per transect and did not produce a proper trend line. They are
Lower Steamboat - ALT (no bankside or middle trees), Lower Fish Creek (no upslope tree), and N Umpqua River Soda
Springs Dam NORTH (no upslope tree).
53
54
Table 3.5. C:N ratios observed in the North Umpqua River basin
55
DISCUSSION
Our results from the foliar isotopic technique suggest that the method may not
be as generally applicable as has been previously suggested. The documentation of
elevated bankside foliar δ15N across all three groups (Salmon, Absent, and Unknown)
and both variable depositional patterns upslope within a National Forest in Oregon
suggest that other biotic and abiotic factors beyond salmon carcass subsidies may be
responsible for the observed foliar δ15N trends (Amundson et al. 2003, D'Amore et al.
2011, Levi et al. 2011, Perakis et al. 2006, Scott et al. 2009, Scott et al. 2008) We are
highly confident in our assumptions that neither Toketee Falls nor Watson Falls are
passable to salmon or other migratory oceanic fishes at levels sufficient to deposit a
marine signal (Appendix E). Yet false positive “salmon-derived” signals (i.e., enriched
δ15N values) were observed at those transects. It is tempting to assume that migratory
salmon were historically present in the Upper Fish Creek watershed (Unknowns)
based upon the relatively enriched foliar δ15N deposition pattern observed. However,
foliar δ15N values were confounded with elevation (e.g. as elevation increased so did
observed foliar δ15N values). Tree location (bankside, middle, upper) was removed as
an explanatory variable from the mixed effects model prevents any analyses about
depositional patterns in the North Umpqua River Basin for any of the groups.
Regression modeling of the soil C:N ratios for Upper Fish Creek indicates
significant positive correlation with the enriched foliar δ15N values observed for the
Unknown transects (Appendices B and D). After accounting for elevation, soil C:N
ratios were still significantly higher for the Fish Creek unknown transects (46.81) than
56
for either of the positive (28.55) or negative (26.98) control groups (r = 0.1937, n=30,
2
p=0.01911). When our foliar C:N values are compared to Koyama (mean = 49.50 and
48.50, for salmon presence and absence, respectively), our relative values are salmon
history dependent (r2= 0.2850, n=29, p=0.237). Unknowns (mean=48.35, n=10) are
equivalent to Koyama, while our salmon presence (mean=40.54, n=13) and absence
(mean=42.52, n=6) are more similar to values reported by Helfield and Naiman
(2001). Sitka spruce (Picea sitchensis) had C:N ratios of 39.21 and 32.73 for reference
and spawning sites, respectively (Helfield and Naiman 2001). If low C:N ratios are
indicative of marine-derived N subsidies, then soils would be a better explanatory
variable than foliar samples because soils are at least one trophic level closer to the
decaying salmon carcasses than the trees. However, that line of reasoning for lower
ratios would also suggest marine subsidies for both the positive and negative control
sites in our study.
Nutrient Retention and N Availability
The foliar δ15N values observed in the North Umpqua Basin are within the
same numerical range and correspond closely to the foliar δ15N range of expected
values in Idaho (+2.75 to -5.50‰). Most of the values were below the relative zero
international standard. Riparian shrubs along the Umpqua River such as blackberry
(Rubus spp.), sedge (Cyperacea spp.), willow (Salix spp.), and cottonwood (Populous
spp.) have reported foliar δ15N values ranging from +0.28 ‰ for the North Umpqua, to
+0.13 ‰ for the South Umpqua, and a higher δ15N value of +2.30 ‰ for the mainstem Umpqua River channel (Coates 2012). Our highest absolute δ15N difference
57
observed between bankside and upslope foliar samples was 1.91 ‰ immediately
downstream of Soda Springs Dam. Several other transects had slightly lower values,
including some serving as negative controls. This is only slightly lower than the 2.3‰
difference reported in central Idaho (Koyama et al 2005) and approximately half of the
values reported at higher latitudes in coastal Washington and Alaska (Helfield and
Naiman 2001, Helfield and Naiman 2002, Hilderbrand et al. 1999). Given the lower
CEC values reported for our study area, which are generally interpreted as indicating
lowered soil fertility and inability to retain nutrients, it seems unlikely that seasonally
available marine nitrogen from salmon carcasses would remain in situ after 60 years of
outright salmon absence above Soda Springs Dam (or impassable natural waterfalls) at
a level comparable to central Idaho after only a 30 year absence. Additionally, the
anticipated effects of observed N-fixers abundance >5% were not observed in our data
set. More than half of our sites had no visible N-fixers present. Linear regression
analyses indicated that type (or species) of N-fixer was important but that the percent
observed was not; the linear mixed effects modeling eliminated the variable
completely from the final model.
Biomass Dependent Subsidies
Current hypotheses regarding the quantity of marine nutrient subsidies from
salmon escapement are generally based upon total biomass deposition on the spawning
grounds. Spawning density (#fish/km) and deposition patterns are also species dependent (Drake and Naiman 2007, Gresh et al. 2000, Lichatowich 1999, Nagasaka
et al. 2006). The North Umpqua River differs from much larger systems such as the
58
Columbia or those in Canada, Alaska, and Russia in terms of total returning spawning
salmon by at least an order of magnitude. However, the North Umpqua River has
several species of salmon returning each year including coho(Oncorhynchus. kisutch),
fall and spring Chinook (O. tshawytscha), summer and winter steelhead, resident
rainbow trout, cutthroat (resident and anadromous) (O. clarkii), and Pacific lamprey
(Entosphenus tridentatus) (Huchko 2012). Cumulatively, a historical, marine nutrient
subsidy to the interior, riparian forests may exist but the observed foliar δ15N
deposition patterns do not reflect that possibility. Nitrogen availability due to isotopic
fractionation and human disturbance may offer more likely explanations for false
“salmon” signals above Toketee Falls and Watson Falls. Human landscape level
alterations are visible via aerial photos and seem to be much more prevalent at the
higher elevations along the mountaintop ridges and alpine meadows of the Southern
Oregon Cascades. Indeed, areas such as Upper Watson, Mowich, and Clearwater have
relatively flat topography compared to the inner valleys of Fish Creek, which makes
them more accessible to human presence and alteration. We believe that our foliar
δ15N sampling does not provide the full story of historical salmonid presence and
absence in the headwaters of the North Umpqua River basin. We think it is only one of
many biotic and abiotic processes which create a confusing landscape level snapshot
of historical, salmonid spawning habitats.
In summary, our study represents a significant re-examination of current
salmon-derived nitrogen signal theory and potential sources of subsidies in interior
riparian forests in SW Oregon. We applied established techniques to provide answers
59
to a serious management question about historical salmonid presence and spatial
distribution. Our goal was to extend the utility and practicality of the technique first
described by Koyama et al (2005). Instead, we have documented serious potential
flaws in the current hypotheses regarding the importance, magnitude, origin, and
expected depositional trends of “marine” nutrient subsidies to interior riparian forests
in SW Oregon. We suggest that alternative explanations and subsidy pathways be
explored in river systems with declining or extinct salmonid populations and
burgeoning human presence.
60
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64
CHAPTER 4: GENERAL DISCUSSION
In November 2012, after 60 years of a barrier, aquatic connectivity was
theoretically restored to the last few kilometers of historical salmonid habitat in the
North Umpqua River basin. As in many other Pacific Northwest rivers, salmon in the
Umpqua Watershed have been in decline for decades, and restoration projects are now
being assessed for effectiveness. Fish ladders and dam removals may solve part of the
problem of habitat accessibility, but the fish themselves must solve the remainder of
the problem through re-colonization. Ultimately each species must adapt to the
restored habitats and ecological challenges.
In Chapter 2, I followed radio-tagged summer steelhead transplanted above
Soda Springs Dam for 1 year to document their behavior and habitat usage. There
were alternative predictions as to how the fish would respond in that newly accessible
habitat. They might move back downstream immediately after release. They might
remain in the area where they were released. They might move upstream into habitat
in Fish Creek. In addition, it was possible that they might spawn somewhere in the
newly accessible habitat. My telemetry data provided unequivocal answers to these
predictions. All of the transplanted steelhead remained within the study area above the
dam. A few fish appear to have left volitionally only after several months spent
holding in faster, deeper flowing areas (Baigún 2003). None of the tagged fish moved
into Steamboat Creek. A small number of the summer steelhead appeared to return to
the base of Soda Springs Dam to spawn, not just hold there until cooler downstream
temperatures return each fall (Dambacher 1991, Holaday 1992, Hostetler 1991). Even
65
though the tagged fish remained above Soda Springs Dam, they did not move further
upstream. Evidence from other dam alteration or removal projects indicates that rates
of re-colonization are highly variable and dependent upon a variety of site-specific
factors (Pess et al. 2012, Pess et al. 2011).
In Chapter 3, I analyzed stable isotope deposition patterns to test the
hypothesis that a historical salmonid nutrient subsidy could be detected spatially in the
headwaters of the North Umpqua River basin. I tested predictions based on accepted
foliar isotopic technique first described in central Idaho (Koyama et al 2005). Those
researchers identified historical salmon spawning habitat above and below barriers
after nearly 30 years of declining adult spawners or extirpated populations. Our results
in the North Umpqua River basin do not support such predictions. Our extensive
background analyses of our data and study area revealed significant differences from
central Idaho, but ultimately there is no simple explanation for our false positive
“salmon” signals above Toketee Falls or the increasing deposition patterns observed
throughout all three salmon history groups. We conclude that human disturbances,
including forest fertilization, spraying of fire suppressing agents, and construction of
roads are the most likely sources of false positive isotopic records.
Future Research
Our research showed that neither telemetry nor stable isotope techniques can
necessarily resolve the important questions for fish passage and habitat restoration.
There can be problems of both false positive and false negative signals from both
these techniques. It seems that answers will be site specific, and empirical studies will
66
probably be required in many cases. We recommend continued observation of recolonization milestones for salmonids passing through the Soda Springs Dam fish
ladder and spawning above the dam. We will need behavioral data for each migratory
species to confirm that passage is possible through the fish ladder and to understand
the specific habitat requirements of each species for holding, spawning, and rearing.
Finally, I recommend collecting non-lethal genetic samples from individual fish
passing through the new Soda Springs Dam fish ladder. That information will be
important to discriminate between wild and hatchery fish and to document their
individual reproductive fitness. Hatchery fish have been documented up to the base of
Soda Springs Dam and could create new challenges for salmonid populations already
stressed by habitat loss (Araki et al. 2008, Araki et al. 2009, Araki and Schmid 2010,
Christie et al. 2011).
Our data provide suggestive clues concerning human disturbance effects on
foliar δ15N riparian deposition, but our dataset and original project scope do not allow
for statistical analyses into these unexpected questions. Potential effects due to the use
of fire retardants in the study area are a definite possibility (Pappa et al. 2008); other
human-related possibilities include successional changes due to timber harvest or fire
suppression (Àgueda et al. 2008, Blank and Zamudio 1998, Durán et al. 2009,
Koyama et al. 2010, Stork et al. 2009, Waples et al. 2008), interrupted hydrological
flows due to roads, as well as documented fertilization on national forest lands (Menke
and Gamble 2012).
67
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76
CHAPTER 6: APPENDICES
77
APPENDIX A: TELEMETRY RESULTS
Fish Salvages
Fish salvages were conducted to rescue and relocate all aquatic species found within
the in-stream work area below Soda Springs Dam. Large groups of people captured
fish, lamprey, freshwater mussels, larval fishes, benthic invertebrates, and
salamanders. Larger animals were relocated in coolers or by hand. Salvaged steelhead
were inspected for physical damage, origin (wild or hatchery), and then dealt with
according to protocol. Animals that could not relocate volitionally or be captured died
onsite. The first two pictures show salvage activities; the third picture shows the
release location upstream of SSD. The fourth picture in the series provides a glimpse
of North Umpqua River riverbed channel completely de-watered. For perspective, the
muddy pool has an estimated depth of less than one meter and the concrete structures
in the riverbed are approximately 6 meters tall. The fifth picture shows relative size
differences between native resident rainbow trout (smaller fish) and migratory summer
steelhead (larger fish); these fish died during salvage efforts in June 2011.
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79
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Radio-tagged Summer Steelhead
Summer steelhead captured via salvage or flyrod were tagged, transported, and then
released upstream of Soda Springs Reservoir at a boat ramp directly across the river
from the Slide Creek Powerhouse. The first two pictures are from the June 2010
salvage, the third picture is a fly-caught fish in October 2010, and the fourth picture is
from a very late arriving summer steelhead salvaged in January 2011.
81
Telemetry Photo showing fish tracked to holding location underneath bedrock bank overhang in fast, deep (≥4m) flowing
water. This type of bedrock ledge or cliff face is abundant within the study area and reflects the typical locations of tagged O.
mykiss in this study.
82
Telemetry results showing areas of incremental tagged fish movements above Soda Springs Dam from June 2010 to May
2011. Majority of tagged fish remained downstream of Fish Creek confluence; 2 fish held at the base of Slide Creek Dam for
several months prior to outmigrating from the study area entirely in late March 2011.
83
USGS Stream Gauge #14315950 graph showing flows recorded during telemetry study. Orange highlighted section indicates
expected spawning behavior timing observed below SSD for summer steelhead (C Street 2009).Red arrows indicate capture,
tag, and release dates for tagged O. mykiss. Green arrow indicates suspected spawning activities for tagged fish; yellow arrow
indicates tagged fish outmigrations from study area (US Geological Survey 2012).
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APPENDIX B: STATISTICAL ANALYSES
Foliar δ15N, C:N ratios, ANOVAS, ANCOVAs, and Regressions
To address prediction 3 in the stable isotope analyses presented in Koyama et al (2005), we assessed several linear regression
models with graphical diagnostics, metrics of data normality, and residual plots. Simple data review was conducted to look for
data independence, normality, kurtosis, variance, and resistance to outliers. We also reviewed the explanatory power (R value)
of each variable, linear model, or ANOVA performed.
85
86
87
88
89
APPENDIX C: MIXED EFFECTS MODELING
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91
92
93
94
95
APPENDIX D: SUBSET TRENDS
Some analyses involving groups separated by observed C/N soils trends, abundance / type of N-fixing species effects, and post
hoc GIS human disturbance data (USFS, R. Menke & K. Gamble, personal communication, 2012) could not be explored fully
in the linear mixed effects model due to small sample sizes.
Salmon ~ C:N Soils residuals (after accounting for Elevation)
R2 = 0.1937, n=30, p=0.01911
96
(p= 0.00436 ** )
C:N Soils ~ Salmon
R2=0.3994, n=30, p= 0.001324 **
97
Salmon ~ C:N Soils (by basin)
R2 = 0.4173, n=30, p=0.04495
98
Salmon ~ C:N Foliar samples (by basin)
R2 = 0.2850, n=29, p=0.237
99
Salmon ~ Dominant N-fixer
R2 = 0.1087, n=46, p=0.02222
100
δ15N ~ Human Disturbance
R2 = 0.1257, n=171, p= 1.953e-06
Was the foliar transect physically bisected by a road? (Yes / No) Half of foliar sites were and half were not disturbed by roads
101
Diamond Lake North Umpqua Fertilized Units received 200 lbs N2 / acre (USFS, R. Menke & K. Gamble 2012)
102
103
APPENDIX E: STABLE ISOTOPE PHOTOS
Waterfalls within North Umpqua River Headwaters
Toketee, Watson, Whitehorse, and Clearwater falls are all impassable to migratory
fish. Watson is located on Watson Creek (tributary to Clearwater River), and
Clearwater Falls is located kilometers upstream of Whitehorse Falls.
Toketee Falls – end of historic anadromy (approx. 37 meters tall)
104
Watson Falls (approx. 83 meters tall)
105
Whitehorse Falls (approx. 6 meters tall)
Clearwater Falls (approx. 9 meters tall)
106
Fish Creek Sub-basin Photos
Fish Creek Confluence with N Umpqua River (Fish Cr = canyon on RIGHT)
Fish Creek Falls - Lower waterfall at 5.1 km (approx. 4.2 m tall), human = 1.8m tall
107
108
109
Subject in picture is 1.8m tall
110
111
112
Foliar Transects and Sample Collection Methods
Pictures indicate the part of the branch collected for foliar analysis (bright green tips
are the current year’s new growth).
113
Subject next to tree is 1.8m tall
113
114
Subject climbing tree is 1.8m tall
115
Subject climbing tree is 1.8m tall
116
117
Subject in tree is 1.8m tall.
118