Upriver Migration and Energetics

• Timing • Patterns • Speed • Energetics • Hell’s Gate

Slide and Fishway

But before we dive into the details, let’s take a moment to consider the athletic performance accomplished by at least some populations…

The Salmon of the Yukon River C.H. Gilbert. 1922. Bulletin U.S. Bureau of Fisheries 38: 317-332.

“As regards the rate at which [Chinook salmon] ascend the river, we have more reliable and complete data for the Yukon than have been secured in any other stream. Records were obtained of their first appearance at a large number of localities. … The first king salmon to reach Dawson in the middle of July, 1920, had been traveling against a consistently rapid current for 29 days, at the rate of 52 miles per day, and during this period, as always within the river, had taken no food.”

The Salmon of the Yukon River C.H. Gilbert. 1922. Bulletin U.S. Bureau of Fisheries 38: 317-332.

“[The chums] entered the river about a week later than the kings, at Tanana they were not more than 10 days behind the latter, and at Dawson they were some 14 days behind the kings. The lower 800 miles of the river, as far as Tanana, were traversed at the rate of 50 miles per day and the next 700 miles, between Tanana and Dawson, were covered at the rate of 35 miles per day.”

Collins et al. 1962 TAFS 91: 1-7

An experimental “endless fishway” was built in which salmon jumped up a series of pools and then slid back down, to test swimming performance to aid design of dams.

In five days a 50 cm sockeye salmon ascended day and night, 2025 m (6648 feet) in 415 circuits of the fishway. It was still climbing when the staff let it go. Not until the fifth day did it show signs of slowing down, and blood lactate levels did not indicate muscular fatigue.

Migration timing between and within species (some patterns and a lot of

variation)

1) Sockeye and Chinook tend to migrate and spawn early

2) Coho tend to migrate and spawn late

3) In Puget Sound pink salmon tend to spawn earlier than chum salmon but farther north this is not the case

4) Chinook and steelhead display extreme variation in migration timing among populations, and also considerable variation within populations (especially steelhead)

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500

1,000

1,500

2,000

2,500

3,000

3,500

8-Jun 8-Jul 7-Aug 6-Sep 6-Oct

Estimated daily counts of adult salmon ascending the Ballard Locks in 2007

sockeye

Chinook coho

Estimated cumulative counts of adult salmon ascending the Ballard Locks in 2007

-

15,000

30,000

45,000

60,000

75,000

8-Jun 8-Jul 7-Aug 6-Sep 6-Oct

Median (50%) dates:

7 July 20 Aug 18 Sept

Monthly entry of chinook salmon as inferred from catches at Rio Vista, including winter, spring, summer,

fall and late fall runs Stone, L. Bull US Fish Comm 16: 203-235.

0

1000

2000

3000

4000

5000

Janu

ary

Februa

ryMarc

hApri

lMay

June Ju

ly

Augus

t

Septem

ber

Octobe

r

Novem

ber

Decem

ber

num

ber o

f sal

mon

cau

ght Sacramento River,

1872

Seasonal changes in the percentage of stream and ocean-type chinook to the total run in the

Fraser River (data from Rich 1942)

Arrival patterns of Chinook salmon destined for parts of the Fraser River basin

Waples, R. S., G. A. Winans, F. M. Utter and C. Mahnken. 1990. Genetic Approaches to the Management of Pacific Salmon. Fisheries 15(5): 19-25.

Migratory timing of Fraser River sockeye salmon populations and pink salmon to the Strait of Juan de Fuca).

Area 20 Date

Early Stuart Early Misc.

Stellako Chilko

Adams/Late

Composite Sockeye

Pinks

Birkenhead

18-Jun 03-Jul 18-Jul 02-Aug 17-Aug 01-Sep 16-Sep

100

200

300

400

500

600

Odd and even year pink salmon can differ in timing: Sashin Creek, Alaska

T.R. Merrell. 1962. In: Symposium on Pink Salmon, N.J. Wilimovsky (ed.)

Perc

ent o

f Tot

al R

un

Mean 50% Date

1 10 20 30 10 20 30 Aug. Sept.

Odd years 1951-1959

Even Years 1950-1958

0

20

40

60

80

Dec Jan Feb March April May June

Ripe

Spent

Counts of upstream migrating (ripe) and spent wild steelhead in two tributaries of the Siuslaw

River, Oregon in 1991-1992 and 1992-1993 (ODFW data, complied by McMillan 2001)

Steelhead are commonly referred to by the season when they return to freshwater (summer or winter) but are more properly referred to by the environment where maturation takes place. Ocean maturing (= winter) steelhead arrive in late winter and spawn shortly thereafter. River maturing (= summer) steelhead arrive in late summer and spawn the following spring.

0

5

10

15

20

25

30

OctDec Feb Apr

Jun

AugOct

Dec Feb

% o

f ann

ual t

otal

winter summer

Entry of adult winter and summer steelhead into the Kalama River, Washington

Winter: mean spawning:

15 April Summer:

mean spawning:

1 Feb

Daily escapement of sockeye salmon into Iliamna Lake and estimated daily total run in 1983

Caveat: Counts in rivers may be affected by fishing…

Changes in fish traits over the run • There are often systematic changes in

the life history traits of salmon over the course of the migration

• Sometimes these reflect differences in timing of discrete populations that differ in those traits

• However, sometimes there are also systematic differences within populations

Older salmon tend to arrive before younger ones (Tuluksak River, Alaska)

0102030405060708090

18-Jun 8-Jul 28-Jul 17-Aug

Sampling date

% a

ge 5

chu

m s

alm

on 1991199219931994

(Molyneaux and DuBois 1998)

Larger salmon of a given age may arrive before smaller ones (Tuluksak River chum)

400

450

500

550

600

650

18-Jun 8-Jul 28-Jul 17-Aug

Sampling date

Mea

n ch

um s

alm

on le

ngth

age 5 malesage 4 females

Male salmon typically precede females (chum salmon)

0

20

40

60

80

100

18-Jun 8-Jul 28-Jul 17-Aug

Sampling date

% m

ales

1991199219931994

Why might older/larger fish and males migrate and arrive before younger/smaller

fish and females?

1) Migratory performance: bigger fish swim faster and so arrive sooner, even though they leave at the same time.

2) Feeding ecology: the optimal duration of time at sea depends on how big you are – smaller fish have a greater need to grow than larger fish.

3) Spawning ground dynamics: smaller fish cannot compete, and so arrive after bigger fish; breeding dynamics favors early arrival by males.

What time of day do salmon migrate upstream?

Primarily at night: kokanee

Lorz and Northcote 1965

Primarily in the day: Sockeye, chinook, steelhead

Columbia River

At night if the water is clear, in the day if turbid: Atlantic salmon

Hellawell et al. 1974

At night in moderate flows, in the day on floods: Atlantic salmon

Potter 1988

Slightly more at night than during the day: Atlantic salmon Brown trout

Allan 1966

What is the relationship between flow and migration?

Increases in flow stimulate migration:

Atlantic salmon Huntsman 1948, Hayes 1953, Saunders 1960, Harriman 1961, Potter 1988, Jonsson et al. 1990

rainbow trout Davies and Sloane 1987

coho salmon van den Berghe and Gross 1989

pink salmon Hunter 1959

What is the relationship between flow and migration?

Increased or high flows retard migration:

Atlantic salmon Alabaster 1970, Hellawell et al. 1974, Jensen et al. 1989

brown trout Stuart 1957, Alabaster 1970, Davies and Sloane 1987

pink salmon Davidson et al. 1943

Relationship between pink salmon migration and flow in Hooknose Creek, AK

August September October November

1952 1200

900

600

300

200

150

100

50

0

Hunter (1959) JFRBC

flow fish

“normalized” fish run

Salmon movement can occur without pulses of stream flow: Skutz Falls, B.C.

Oct.21 22 23 24 25 26 27 28 29 Nov. 5 6

1400 1200 1000 800 600 400 200

25 20 15 10 5

Neave (1943) JFRBC

flow

fish

Upriver travel can be very rapid

• Chinook – Yukon River = 36 km/day (radio tagging) – Yukon tag/recovery ~ 40 to 77 km/day – Columbia River: 1134 km at 26 km/day

(pre-dam)

Chum - Amur River: 1193 km at 45 km/day

Jumping ability Steelhead > Chinook > coho > chum = pink

Reiser et al. 2006 TAFS

Energetics of Fraser River Sockeye

Components of Research 1. Estimate travel rate (tagging, counts) 2. Estimate rate of energy consumption

as a function of activity (lab) 3. Determine rate of energy depletion

during migration 4. Estimate actual swimming speed

Energetics of Fraser River Sockeye

Analogy: Car 1. Determine Seattle to Olympia

distance and time of travel 2. Determine the fuel consumption rate 3. Measure the gas used 4. Estimate the driving pattern

Takla Lake

Stuart Lake Francois Lake

Quesnel Lake Horsefly River

Chilko Lake Adams River

Shuswap Lake

Hell’s Gate

Fraser River system, B.C.

Speed of upriver travel • Sockeye

o Based on run peaks at different sites: oStuart, Bowron, Horsefly = 48 km/day oChilko and Stellako = 34 km/day oAdams = 27 km/day

o Stuart Lake: Lummi Island to Forfar Creek:

o1152 km in 27 days = 43 km/day (tagging)

Relationship between percent body fat and migratory distance

Gilhousen 1980

0

4

8

12

16

0 250 500 750 1000 1250

Distance upriver (km)

perc

ent b

ody

fat

Early StuartChilkoAdams

Energy remaining (% of initial) for Fraser River sockeye from entry to

death on spawning grounds Race Sex Fat Protein Distance

(km) Elevation

(m)

Stuart M 9 64 1152 693 F 5 45

Chilko M 17 61 725 1170 F 7 41

Adams M 15 70 483 347 F 13 61

Gilhousen. 1980. IPSFC Bull 22

Populations and species of salmon with more arduous migrations have more fat

when they leave the ocean

0

3

6

9

12

15

18

1 10 100 1000 10000 100000 1E+06

distance (km) x elevation (m)

% fa

t at r

iver

ent

ry

Data sources: Brett 1995, Mathisen et al. 1988, Williams et al. 1986, Kizevetter 1948, Davidson and Shostrom 1936, etc.

Sockeye salmon were placed in a swimming tunnel, water velocity and temperature were controlled, body size was measured, and oxygen consumption and energy use were recorded after determining the swimming capacity of the fish.

Energetics of migrating sockeye

0

5

10

15

20

25

30

35

0 1 2 3 4 5 6 7Speed (km/hr)

Ener

gy c

ost (

kcal

/kg/

km)

Optimum

Observed in coastal waters

Estimatedduringupriver

migration

Maximumsustained

Burst

Observed duringupriver

migration

For a 2.27 kg, 61 cm sockeye least energetic cost is 1.8 km/h. River Speed: equivalent to 4.3 km/h

Brett, J.R. 1983. In: Aspey and Lustick (Eds.) The cost of survival in vertebrates.

Fraser River sockeye salmon catch

05

101520253035

1896 1904 1912 1920 1928 1936 1944 1952 1960 1968 1976 1984 1992 2000

Year

mill

ions

of s

ocke

ye

Note: 4-year cycle, and drop in catch after 1913

Fraser River at Hell's Gate, 1897, looking upstream before the railway was built on the East Bank

At Hell’s Gate the river is so constricted that it varies by nearly 30 m during a year

Jackson. 1950. IPSFC Bull. III

Hell’s Gate • The Fraser River was probably always

hard for salmon to ascend at Hell’s Gate (and other locations).

• 1913 - dumping of rock material from railway construction virtually blocked passage of fish; millions died unspawned

• 1914 Feb. 23: a huge rockslide, caused by Canadian Northern Railway tunneling, again greatly restricted flows

Sockeye salmon blocked at Hell's Gate, 1913 www.saxvik.ca

Hell's Gate, February, 1914, after the collapse of the rock cliff above the Great Northern Railway Tunnel

Hell’s Gate: 1915 • After the slide it was virtually impassable to

fish from levels 25 to 40 feet and difficult from 10 to 60.

• Vertical drop: 9 feet (after rocks removed). • Velocity: 5.0 to 6.75 meters per second

Hell’s Gate, 1915, excavated rock from slide on east bank deposited onto rock shelf on west bank.

Hell's Gate, 1914, looking upstream, temporary fishway built on the east bank to aid adult salmon migrating up-river

www.saxvik.ca

Hell’s Gate

• Fishways now designed to pass 26,000 fish per hour in daylight. At the peak of the migrations they may pass over 500 salmon per minute.

• Aid passage at 10 to 70 feet elevations.

Correlation between mortality of sockeye salmon and temperature

(Gilhousen 1990)

R2 = 0.49

0

10

20

30

40

50

10 12 14 16 18

Spawning ground temperatureP

erce

nt m

orta

lity

Raft River

R2 = 0.36

0

10

20

30

12 14 16 18

Temperature at Hell's Gate

Perc

ent m

orta

lity

Early Stuart Lake

r2 = 0.28

r2 = 0.21

12

14

16

18

20

22

24

1950 1960 1970 1980 1990 2000 2010

ColumbiaFraser

Water temperatures (e.g., August means) have been getting warmer…

Med

ian

mig

ratio

n da

te

1940 1960 1980 2000

175

180

185

190

r2=0.43, p<0.0001

One response by salmon has been to avoid the warmer water. Columbia River sockeye are migrating earlier than in the past; steelhead are migrating later.

Future effects of climate change?

Estimated swimming speeds of migrating sockeye salmon (Hinch, Quinn data)

0

5

10

15

20

25

20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 >300

swimming speed (cm/sec)

% fr

eque

ncy

coastalriver: constrictedriver: unconstricted

150 cm/sec = 5.4 km/h

or 2.5 body lengths/sec for a 60 cm fish

Year

Mea

n Ju

ly T

empe

ratu

re (o

C)

1950 1960 1970 1980 1990 2000

1718

1920

21

r2=0.42, p<0.001

Temperatures (e.g., the Columbia River at Bonneville Dam) have been getting warmer

Hell’s Gate Cross section at reference line station 20

Primary Sockeye Migration

Left Bank

Downstream view

Dep

th in

Fee

t

100 50 0 -50 -100

Jackson. 1950. IPSFC Bull. III

Max: 61 feet

Min: 12 feet

1912 to 1948

Max: 93 ft

Min: 8 ft

Steelhead and chinook salmon counts at the fish ladder as a function of time of day: Lower

Granite Dam, Snake River, in 1992.

0 2 4 6 8 10 12 14 16 18 20 22

0 2 4 6 8 10 12 14 16 18 20 22

2 4

6 8

10

Mid

-day

4 8 12 16 20 24 Hour of the day

Hayes (1953) B.F.R.B.C.

Timing of Atlantic salmon ascending the Le Havre River, Nova Scotia