Salmonid Feeding and Growth at Sea 1. Habitats and ...courses.washington.edu/fish450/Lecture...
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Salmonid Feeding and Growth at Sea
1. Habitats and communities of salmon 2. Diet of salmon 3. Patterns of feeding 4. Growth 5. Density and temperature
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Feeding and growth of salmon at sea
• Salmon time their seaward migration to take advantage of marine prey
• Salmon grow rapidly and achieve > 99% of their total weight at sea
• Salmon diets reflect their size and species, and the prey available to them
• Growing conditions vary among years, with ocean conditions and salmon density
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Salmon in the central North Pacific and Bering Sea
Asia
Alaska
British Columbia
The distributions are large, and overlap broadly between species and populations.
What is the nature of the region that they occupy?
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North Pacific Ocean current systems (Dodimead et al. 1963)
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Approximate area of oceanic domains and prevailing current directions in the Northeast Pacific Ocean (Ware and McFarlane 1989)
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0%
10%
20%
30%
40%
50%
60%
70%
55 54 53 52 51 50 49 48 47 45 44 44 42 41 39 38 36
Latitude
Perc
ent o
f Tot
alCatches of fishes and squid in gill nets along
155o W by the Oshoro Maru, July 1984
n = 3999
n = 2897
n = 694
n = 199 Salmonids
Pacific Pomfret
Blue Shark, Albacore, Ommastrephes bartami Tuna, billfish, Mackerel, mahi mahi, Yellowtail
Open ocean communities
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0
5
10
15
20
25
30
herrin
g
salm
onids
anch
ovies
stick
lebac
ks
shark
soth
ersq
uid
sand
lanc
e
atheri
nids
smelt
gadid
s
rockfi
shjac
ks
coastalinshore
0
10
20
30
40
50
60
70
herrin
g
salm
onids
anch
ovies
stick
lebac
ks
shark
soth
ersq
uid
sand
lanc
e
atheri
nids
smelt
gadid
s
rockfi
shjac
ks
coastal
inshore
California Current: dominated by herring
Alaska Current: dominated by salmon
Orsi et al. 2007 AFS symposium 57
% of fishes caught, by family, in two current systems
Coastal and inshore communities > 70%
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Thysanoessa longipes
15 – 23 mm
Themisto pacifica
2 – 8 mm
Common crustaceans eaten by salmon
Euphausiids (krill) Hyperiid Amphipods
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Some fishes eaten by salmon at sea
Leuroglossus schmidti ~135 mm SL
Stenobrachius leucopsarus ~90 mm SL
Pleurogrammus monopterygius ~40 mm SL Atka mackerel
Hemilepidotus spp. ~15 mm SL
Northern lampfish (lanternfish) Northern smoothtongue - Bathylagidae
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Fishes eaten by salmon (Puget Sound)
Pacific herring
Northern anchovy
Sand lance
Surf smelt
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Limacina helicina
2 – 3 mm
Berryteuthis anonychus
Gonatopsis borealis
Pteropods Squid
Mollusks eaten by salmon
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Gelatinous zooplankton: ctenophores, medusae, salps (tunicates)
Beroe spp. ~45 mm TL
Salpa sp. ~20 mm TL
Aglantha digitale ~15 mm TL
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Energy density of salmon prey organisms
0 500 1000 1500 2000 2500
Stenobrachius leucopsarus (112 mm SL)
Stenobrachius leucopsarus (64 mm SL)
Berryteuthis anonychus (82 mm ML)
Berryteuthis anonychus (86 mm ML)
Hemilepidotus sp. (21 mm SL)
Thysanoessa longipes (23 mm TL)
Tarletonbeania crenularis (43 mm SL)
Thysanoessa longipes (19 mm TL)
Parathemisto pacifica (Females w/ young)
Limacina helicina (3 mm TL)
Hippoglossus stenolepis (19 mm SL)
Squid (22 mm ML)
Sebastes sp. (11 mm SL)
Clione limacina (16 mm TL)
Salpa sp. (35 mm TL)
500 1000 1500 2000 2500 0
Salpa, Beroe
pteropods, Themisto; juv. fish, euphausiids juv. squid
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Annual cycle of average zooplankton density at Ocean Station P
050
100150200250300
Jan
Feb
Mar
April
May
June Ju
lyAu
gSe
pt Oct
Nov
DecZo
opla
nkto
n (g
/100
0 m
3)
Brodeur et al. (1996)
(mean and standard error)
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Synchronous downstream migration of pink salmon fry and density of zooplankton in Prince
William Sound, Alaska (Cooney et al. 1995).
0
1
2
3
4
5
21-Mar 20-Apr 20-May 19-Jun
Zoop
lank
ton
(ml/m
3)
0
20
40
60
80
100
Pink
sal
mon
(tho
usan
ds)zooplankton
pink salmon
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Wakatake maru
Food habits of salmon vary among species, years, and among fish of different sizes
Nancy Davis
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0%
20%
40%
60%
80%
100%
91 92 93 94 95 96 97 98 99 00 01 02 03
UnidOtherGelatinousFish PteropodSquidCrab larvAmphipodCopepodEuphausiid
0%
20%
40%
60%
80%
100%
91 92 93 94 95 96 97 98 99 00 01 02 03
UnidOtherGelatinousFish PteropodSquidCrab larvAmphipodCopepodEuphausiid
Salmon prey composition Bering Sea Basin, summer 1991-2003
Coho
Chinook
no data no data
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0%
20%
40%
60%
80%
100%
91 92 93 94 95 96 97 98 99 00 01 02 03
UnidOtherGelatinousFish PteropodSquidCrab larvAmphipodCopepodEuphausiid
0%
20%
40%
60%
80%
100%
91 92 93 94 95 96 97 98 99 00 01 02 03
UnidOtherGelatinousFish PteropodSquidCrab larvAmphipodCopepodEuphausiid
0%
20%
40%
60%
80%
100%
91 92 93 94 95 96 97 98 99 00 01 02 03
UnidOtherGelatinousFish PteropodSquidCrab larvAmphipodCopepodEuphausiid
Sockeye
Pink
Chum
Prey of salmon collected from the Bering Sea basin 1991 to 2003
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sockeye chum pink chinooksockeye 52 88 54chum 52 26pink 55chinook
Diet overlap Bering Sea basin, summer 1991 - 2003
LowModerateHighVery high 75 - 100%
0 - 24% 25 - 49%
50 - 74%
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0
25
50
75
100
0 100 200 300 400
cohoChinook
Body length (mm)
% fi
sh in
the
diet
Ontogenic diet shift: coho and Chinook salmon consume
a higher proportion of fish as they grow
Daly et al. 2009
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Diet vs. body size
Body Weight (g)
0%20%40%60%80%
100%
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Time of day
0
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30
40
0600-0800
0900-1100
1200-1400
1500-1700
1800-2000
2100-2300
0000-0200
0300-0500
Euphausiids and copepods
Sockeye: diel change in prey composition
Prey
wt (
g)
Euphausids and copepods
0
10
20
30
40
0600-0800
0900-1100
1200-1400
1500-1700
1800-2000
2100-2300
0000-0200
0300-0500
Time of day
Fish Fish
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Data storage tags record temperature and pressure (depth)
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Attachment of the DST to salmon
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Chum salmon
TD Tag 1565 - Hokkaido Chum Salmon
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6
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9 16 21 25 28 1 4 8 11 15 18 22 25 29 1 5 8 12 15 19 22August
Tem
pera
ture
( C)
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50
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150
200
250
300
350
400
Dept
h (m
)
July September
Tagged: 7/9/02Location: 56-30N, 179-00ELength: 618 mm, Age: 0.3
Recovered: 9/24/02Nemuro coast, Hokkaido, JapanLength: 655 mm, Weight: 3.4 kg
= night
recuperation period coastal period
diel vertical migration
Depth
Temperature
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Growth of salmon at sea
Pink Chum Chinook Sockeye Coho Steelhead Masu
Smolt weight (g)
0.22 0.4 5 to 18 10 18 50 18
Adult weight
(kg)
1.63 3.73 7.22 2.69 3.02 3.48 1.45
Full years at sea
1 2,3,4 2,3,4 2,3 1 2 1
Data from McGurk 1996, Groot and Margolis 1991, Bigler et al. 1996, Burgner et al. 1992, Kato 1991.
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0
5
10
15
20
25
370 390 410 430 450 470 490 510 530Length (mm, MEH)
femalesmales
In many cases, male salmon grow faster than females (i.e., are longer at a given age)
4-yr old sockeye salmon from Woody Island, Iliamna Lake
Perc
ent o
f fus
h m
easu
red
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Males may also be heavier for a given length
Woody Island sockeye salmon
0
1
2
3
4
5
350 400 450 500 550 600
Males
Females Mas
s (k
g)
Length (MEH, in mm)
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Factors affecting the growth of salmon at sea
• Density (intra-specific competition) • Ocean conditions
– Productivity – Temperature
• Inter-specific competition
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600
640
680
720
760
0 15 30 45 60Chum salmon abundance (millions)
Fork
leng
th (m
m) males age 4
females age 4
Mean fork length of Hokkaido chum salmon as a function of the number of adult chum salmon (catch plus escapement)
that year (from Kaeriyama 1998).
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Mean length (males and females combined, for fish spending two years at sea) of adult Bristol Bay sockeye salmon as a
function of the number of adults returning to spawn that year.
480
490
500
510
520
530
540
0 20 40 60 80Millions of adult salmon
Mea
n le
ngth
(mm
) 1958-761977-971998-02
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Hypothesized effects of ocean temperature and density on sockeye salmon growth rate
Abundance
Body Size at Age
Temperature 1
Temperature 2
(Peterman 1984)
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photo by Richard Bell
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Growth History
Scale Radius
Fish
Len
gth Individual circuli
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Back-calculated lengths of chinook salmon (in mm) at different ages, as a function of their final age at maturity
(data from Parker and Larkin 1959).
AGE Age at maturity
Sample 2 3 4 5 6
2 4 266 3 27 220 508 4 150 183 466 676 5 60 171 425 622 813 6 8 122 368 549 730 917
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…the end
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0
5
10
15
20
91 92 93 94 95 96 97 98 99 00
SockeyeChumPinkChinookTotal
Central Bering Sea Mean CPUE per tan 1991-2000
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Age notation: 1 . 2
No. freshwater winters No. ocean winters
Species Common ages & maturity in catch
Sockeye 1.1, 2.1, 1.2, 2.2 immature, maturing
Chum 0.1, 0.2, 0.3 immature, maturing
Pink All maturing 0.1
Coho All maturing 1.1, 2.1
Chinook 1.1, 1.2, 1.3 immature
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Stomach contents for salmon sampled in the Strait of Juan de Fuca
0 4
0
80
< 40 cm 40-60 cm Total
Coho
0 4
0
80
Chinook % s
tom
ach
volu
me
< 40 cm 40-60 cm
> 60 cm
> 60 cm
Fish Crustaceans Miscellaneous
Total
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Stomach contents of salmon in the Strait of Juan de Fuca
0 4
0
80
< 55 cm >55 cm Total
Sockeye
0 4
0
80
Pink % s
tom
ach
volu
me
< 55 cm < 55 cm
Fish Crustaceans Miscellaneous
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Biomass of Salmonids
0200400600
Coastal shelf Coastaldownwelling
Coastalupwelling
Domain
Biom
ass
(T x
100
0)
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LowModerateHighVery high 75 - 100%
0 - 24% 25 - 49% 50 - 74%
sockeye chum chinooksockeye 49 21chum 28
chinook
sockeye chumsockeye 80chum
Summer, basin, even yrs 1992-2002
chum chinookchum 30
chinook
Fall, basin, 2002
Fall, Aleutian Is., 2002
Fall, EBS shelf, 2002
Sockeye N=78 Chum N=360 Chinook N=84
Sockeye N=32 Chum N=58
Chum N=54 Chinook N=16
Comparison of Diet Overlap-Summer and Fall
sockeye chum pink chinooksockeye 56 76 57chum 58 26pink 45chinook
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sockeye chum pink chinooksockeye 52 88 54chum 52 26pink 55chinook
sockeye chum pink chinooksockeye 56 76 57chum 58 26pink 45chinook
Sockeye Chum Pink ChinookSockeye 44 83 54Chum 40 28Pink 63Chinook
All years, 1991-2003
Even years, 1992-2002
Odd years, 1991-2003
Diet overlap Bering Sea basin, summer
Sockeye N=724 Chum N=1523 Pink N=1331 Chinook N=193
Sockeye N=577 Chum N=1413 Pink N=156 Chinook N=274
LowModerateHighVery high 75 - 100%
0 - 24% 25 - 49% 50 - 74%
(low pink)
(high pink)
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Relationship between the size of salmonids and the size of their prey
Keeley and Grant 2001. CJFAS
1
10
100
1000
1 10 100 1000Fish length (cm)
Pre
y le
ngth
(mm
)
Fishes (all environments combined)
Ocean invertebrates
Freshwater invertebrates (streams and lakes combined)
Keeley and Grant (2001)
Salmonid Feeding and Growth at SeaFeeding and growth of salmon at seaSlide Number 3Slide Number 4North Pacific Ocean current systems (Dodimead et al. 1963)Slide Number 6Slide Number 7Catches of fishes and squid in gill nets along 155o W by the Oshoro Maru, July 1984Slide Number 9Slide Number 10Slide Number 11Slide Number 12Slide Number 13Slide Number 14Slide Number 15Annual cycle of average zooplankton density at Ocean Station PSynchronous downstream migration of pink salmon fry and density of zooplankton in Prince William Sound, Alaska (Cooney et al. 1995).Slide Number 18Slide Number 19Slide Number 20Slide Number 21Slide Number 22Slide Number 23Slide Number 24Slide Number 25Slide Number 26Slide Number 27Growth of salmon at sea Slide Number 29Slide Number 30Factors affecting the growth of salmon at seaMean fork length of Hokkaido chum salmon as a function of the number of adult chum salmon (catch plus escapement) that year (from Kaeriyama 1998).Mean length (males and females combined, for fish spending two years at sea) of adult Bristol Bay sockeye salmon as a function of the number of adults returning to spawn that year. Hypothesized effects of ocean temperature and density on sockeye salmon growth rateSlide Number 35Slide Number 36Back-calculated lengths of chinook salmon (in mm) at different ages, as a function of their final age at maturity (data from Parker and Larkin 1959).Slide Number 38Slide Number 39Slide Number 40Stomach contents for salmon sampled in the Strait of Juan de FucaStomach contents of salmon in the Strait of Juan de FucaSlide Number 43Slide Number 44Slide Number 45Slide Number 46Relationship between the size of salmonids and the size of their prey