Pattern and process in upstream migration of steelhead in the Columbia and Snake rivers
Christopher C. CaudillMatt Keefer
Department of Fish and Wildlife SciencesFish Ecology Research Laboratory
UI Fish Ecology Research LabUI Project PI: Ted Bjornn (late 1990’s – 2001)Chris Peery (2001‐2008)Staff Scientists (MSc)Charles BoggsTami ClaboughJeff GarnettMike JepsonMatt KeeferGeorge NaughtonSam BourretChris NoyesEric JohnsonHattie Zobbot
Post‐docs:Tracy BowermanGeoff Moret
Graduate StudentsMatt DunkleCharlie ErdmanMark KirkChanning Syms
Full Time Technicians & StaffTim BlubaughTravis DickKal JohnsonDan JoostenMatt KnoffLes LayngMark Morasch
Seasonal and TemporaryTheresa TillsonGrant BrinkKate AbbottMike TurnerNoah HubbardKaan OralRobert Escobar
UI Fish and Wildlife SciencesBrian Kennedy
UI Center for Ecohydraulics ResearchRalph BudwigDaniele Tonina
NOAA‐FisheriesMary Moser Brian BurkeKinsey FrickLisa CrozierPaul Chittaro
OSUCarl SchreckMike KentMike ColvinJim Peterson
UC DavisFrank LogeDon ThompsonPeter Green
CRITFCSean NarumJon HessBrian McIlraith
ODFWCam SharpeTom Friesen
Blue Leaf EnvironmentalMark TimkoLeah Sullivan
USACESean TackleyBob WertheimerDave GriffithRich PiaskowskiDerek FryerSteve JuhnkeChris PinneyEric Hockersmith
Aims• Pattern and process in steelhead upstream migration behaviors:– Route finding and orientation (Chinook salmon as model)– Overshoot and fallback– Straying– Behavioral thermoregulation– Mainstem survival and overwintering– Transport effects– (Not: Kelting, iteroparity, timing, dam passage etc.)
• Terminology• Uncertainties and areas of concern
Overshoot scale 1‐100s kmPrecision of natal site fidelity?
‐mechanisms of site fidelity?‐mechanisms contributing to imprecision?
Outmigrant B
Outmigrant A
Sequential Imprinting Hypothesis
● Novel odors● Elevated stress
● Environmental and ecological gradients
● Incubation ● Rearing● Smoltification
● Incubation ● Rearing● Smoltification
Homing
●Migration stimulates imprinting
● Long‐distance or complex
migrations may require more ‘waypoints’
● Critical neural development and
memory formation
Harden‐Jones (1968)Brannon (1982)
Mechanism: insufficient imprinting
N D J F M A M J J A S O N D J F M A M J J
Thyr
oxin
e le
vel
HatcheryWild
Imprinting threshold
Hatching-emergence Rapid growth Parr-smolt transformation
Modified from Dittman & Quinn (1996)
● Hatchery fish stray more: lower juv hormone levels, reduced neural development, retarded imprinting, etc.
Endocrine control of imprinting
Straying mechanisms• Juvenile effects
– Olfactory imprinting, sequential imprinting, neural development, endocrine physiology
– Hatchery rearing• Adult effects
– Olfactory recognition, sensory failure, memory, age– Reproductive behaviors
• Environmental / Landscape effects– Thermal refuge; Catastrophe
Straying lexicon• Permanent straying
– Some don’t breed with recipient pop: loss from donor population, possible colonist
– Breeders: donor loss + recipient gain
Donor population
Recipientpopulation
● ‘Undesirable’ breeding stray = recipient population
risk
Colonist
Straying lexicon• Temporary straying
– ‘Wandering’ ‘Exploring’ ‘Proving, Testing’ ‘Thermoregulating’
– Just a detour: Exit before breeding
Donor population
● Differentiating permanentand temporary straying is often challenging due to ambiguous outcomes
The lexicon: how to estimate straying?• Spatial scale is critical, yet difficult to define
– Genetic distance more relevant than geographic distance; Hatchery practices confound
• All potential straying locations (and times) are rarely surveyed for donor populations
• All fish origins are rarely known for recipient populations
• Ambiguous outcomes are common
● Despite large straying literature, comparisons are challenging
Mainstem Route‐finding (Chinook salmon)
Keefer, M. L., C. C. Caudill, C. A. Peery, and T. C. Bjornn. 2006. Route selection in a large river during the homing migration of Chinook salmon (Oncorhynchustshawytscha). Canadian Journal of Fisheries and Aquatic Sciences 63:1752‐1762.
Overshoot and downstream movements (Chinook salmon)
OvershootUndershoot Males, Hatchery
Keefer, M. L., C. A. Peery, and C. C. Caudill. 2006. Long‐distance downstream movements by homing adult chinook salmon. Journal of Fish Biology 68:944‐950.
Keefer, M. L., C. C. Caudill, C. A. Peery, and C. T. Boggs. 2008. Non‐direct homing behaviours by adult Chinook salmon in a large, multi‐stock river system. Journal of Fish Biology 72:27‐44.
FallbackInflates dam counts.
Delay, injury, mortality.
Average reduction in Prob(escape) of ~0.133for steelhead that fellback ≥ 1 time(s)
Keefer, M. L., C. A. Peery, W. R. Daigle, M. A. Jepson, S. R. Lee, C. T. Boggs, K. R. Tolotti, and B. J. Burke. 2005. Escapement, harvest, and unknown loss of radio‐tagged adult salmonids in the Columbia River ‐ Snake River hydrosystem. Canadian Journal of Fisheries and Aquatic Sciences 62:930‐949.
Steelhead
30 May 19 Jun 9 Jul 29 Jul 18 Aug 7 Sep 27 Sep 7 Oct
Tem
pera
ture
(10
year
mea
n)
14
16
18
20
22
Cou
nt @
Bon
nevi
lle D
am (1
0 ye
ar m
ean)
0
2000
4000
6000
8000
10000
12000
~Migration constrained~
Temporary straying & behavioral thermoregulation
Sockeye salmon
Fall Chinook
Summer Chinook
~Optimal temps~
~Optimal temps~
1-Apr 21-Apr 12-May 2-Jun 23-Jun 14-Jul 3-Aug 24-Aug 14-Sep 5-Oct 26-Oct
Tem
pera
ture
(C)
06
8
10
12
14
16
18
20
22
24
John Day Pool Deschutes River
Mouth
John Day Pool
John Day Ladder
John Day Ladder
John Day Pool
John Day Ladder
John Day Ladder
Bonneville Pool
The Dalles Pool
Bonneville Tailrace
Deschutes River Mouth
Summer Chinook Salmon start date
FallChinook Salmonstart date
Behavioral thermoregulation
Spring, summer & fall Chinook
● Chinook salmon migrate more directly, userefugia less (median = 3 d for fall Chinook):less scope for thermoregulation
1-Apr 21-Apr 12-May 2-Jun 23-Jun 14-Jul 3-Aug 24-Aug 14-Sep 5-Oct 26-Oct
Tem
pera
ture
(C)
06
8
10
12
14
16
18
20
22
24
Little White Salmon River
White Salmon River
John Day PoolJohn Day Ladder
John Day Ladder
LW River
Little White Salmon River
Deschutes River/Mouth
John Day Ladder
Bonneville Pool
Klickitat River
Little White Salmon River
White Salmon River
Little White Salmon RiverDeschutes River
John Day River
Behavioral thermoregulation
● Steelhead migration relatively flexible andrefugia use often lasts weeks
Summer steelhead with archival tags
Caudill et al. in prep
Behavioral thermoregulation
1 Jun 1 Jul 1 Aug 1 Sep 1 Oct 1 NovMed
ian
mig
ratio
n ra
te (k
m*d
-1)
0
10
20
30
40
50
60
70
1996199720002001
Summer steelhead● Upstream migration ratesfrom radiotelemetry studiesshow reduced swim speeds,long holding behaviors
Keefer et al. 2004
Percent of population that used cool-water tributaries (%)
20 30 40 50 60 70 80 90
Med
ian
pass
age
time
(d)
0
10
20
30
40
50
60
CWRTUC
SNK
WWR
LFH
GRRYAK
UCR
UMA
HAN
SAL
JDRIMR
Used thermal refugiaDid not use refugia
BON TDA
JDAMCN
IHR
PRD
Snake R.Columbia R.
Frying PanFire!
SpawningFW
Rearing
MarineGrowth
Juvenile Outmigration (Spring)
Adult Spawning Migration
Barge (fast)
In‐river (slow)
Transportation and permanent straying
Does Juvenile Experience Affect Adult Migration and Homing?
Year2002 2004 2006 2008 2010
Pro
porti
on T
rans
porte
d
0.0
0.2
0.4
0.6
0.8
1.0
1.2
Large proportions of SNR steelhead and Chinook salmon are transported annually
SpawningFW
Rearing
MarineGrowth
Juvenile Outmigration (Spring)
Adult Spawning Migration
Barge (fast)
In-river (slow)
Snake River: Two Migration Corridors
Interrogate for PIT tag
PIT tag
Juvenile Barging Affects Adults
Homed Strayed Unknown Fell back Homed Strayed Unknown Fell back
Mea
n pe
rcen
t (%
+/-
1 SE
)
0
10
20
3060
70
80
90
100
In-riverBarged
A) Chinook Salmon B) Steelhead
*
*
*
*
*
* ***
Keefer et al. 2008
Mechanism: adult behaviors
Keefer et al. (2008); also see Chapman et al. (1997) Number of fallback events
1 2 3 4 5 6 7 8 9 10
Perc
ent (
%)
0
2
4
6
8
10Pe
rcen
t (%
)0
2
4
6
8
10
In-riverBarged
A) Chinook salmon
B) Steelhead
Barged Chinook salmon:~3.8 times more likely to fall back at dams (P<0.001)
Barged steelhead:~1.7 times more likely to
fall back (P<0.023)
Orientation / Navigation problems!
FCRPS Overwintering RT Study (ongoing)
Keefer et al. 2014 Steelhead overwintering letter report
“Bypass” or “overshoot” or “temporary straying” or ??
• Additionally “Bypass” is frequently used in literature for downstream passage routes
• “Overshoot” accurate in most cases• “Temporary straying” or “permanent straying” cover other
situations (e.g., “undershoot”, behavioral thermoregulation)• “Wandering”, “testing”, etc. not recommended because do
not relate to a known mechanism or to outcome• Recommend use of explicit operational definitions and criteria
http://www.merriam‐webster.com/dictionary/bypass
Steelhead overshoot, homing and straying
• Permanent and temporary straying are challenging to distinguish
• Permanent straying is integral to metapopulation dynamics (i.e., not all straying is bad)
• Overshoot and temporary straying are integral to homing process in steelhead
• Adults barged as juveniles have higher rates of straying, undershoot, (overshoot?), fallback, unsuccessful homing
• Fallback associated with increased mortality• Improved downstream routes would benefit overshoots and kelts
Questions?