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Ocean life
Kees Camphuysen Royal Netherlands Institute for Sea Research, Texel
Theatrum orbis terrarum, Abraham Ortelius, 1590
Giant oarfish Regalecus glesne
Steller’s Sea Cow Hydrodamalis gigas, weighing ± 10 tons or more, ate mostly kelp. became known to science in 1741, inhabited kelp beds of the Commander Islands,
W Bering Sea.
Its meat was considered “as good as the best cut beef”, which led to its extermination at the hands of whalers.
Species was slaughtered to extinction soon after its discovery, the last known live individual was taken in
1768.
Steller’s Sea Cow Hydrodamalis gigas,
1741-1768
• discovered 1741, Bering Sea
• no pelt, no tusks, non-aggressive
• very tasty meat (beef), easy to ‘harvest’
• extinct 1768
Castro & Huber 1997. Marine Biology. WC Brown Publishers
© CJ Camphuysen Northern Gannet
© CJ Camphuysen Sooty Shearwater
8
storm-petrels 32cm span, 30g mass (Halocyptena microsoma)
White-faced Storm-petrel
Blue Whale 33m length max, 190,000kg (Balaenoptera musculus)
Magnificient Frigatebird
Gentoo Penguin
© CJ Camphuysen Southern Elephant Seal
© CJ Camphuysen Great Black-backed Gulls
Contents
• habitats & resources
• scale & transitions
• foraging techniques
• foraging interactions
towards a deeperunderstanding of distribution patterns of marine apex predators
charismatic megafauna
10m high North Sea waves
Marine resources
Phytoplankton
Benthic invertebrates
Zooplankton
Small fish
Squids
Predatory fish
whales
dolphins
seals
seabirds
Trophic level 1 2 3 4 5 6 7
1
2 3
1
2
piscivorous fish and dolphins (salmon, sharks, dolphins)
4
8
5
4
planktivorous whales (baleen whales)
3
19
5-6 64-5Food webs
Open oceans
Continental shelves
Upwelling regions
Bathypelagic
21
The basic ecological divisions of the ocean
BENTHIC ENVIRONMENTS(benthos)
PELAGIC ENVIRONMENTS(plankton & nekton)
0m
200m
1000m
2-3000m
4000m
6000m
10000m
high waterlow water
littoral sublittoralbathyal
abyssal
hadal
Neritic Oceanic
Hadalpelagic
Epipelagic
Mesopelagic
Abyssopelagic
Shelfbreak
supralittoral
Glossary:plankton = passive against currentsnekton = active against currentsbenthos = sessile
22
Strictly areal
Surface feeders
Shallow divers
Deep divers
no water contact
not underwater or shallow
plunging, <1.5m
deepplunging, pursuit
plunging, pursuit diving
<25m
pursuitplunging,
pursuit diving<250m
pursuit diving<500m
1600m
1775m
2130m2250m
3000m
Shelf break
Dive depth
23
BENTHIC ENVIRONMENTS(benthos)
0m
200m
1000m
2-3000m
4000m
6000m
littoral sublittoralbathyal
abyssal
hadal
Neritic Oceanic
Hadalpelagic
Epipelagic
Mesopelagic
Bathypelagic
Abyssopelagic
Shelfbreak
shallow water species
shelf & oceanic ecotypes
shelf break assembly
deep diving, oceanic taxa
24
BENTHIC ENVIRONMENTS(benthos)
0m
200m
1000m
2-3000m
4000m
6000m
littoral sublittoralbathyal
abyssal
hadal
Neritic Oceanic
Hadalpelagic
Epipelagic
Mesopelagic
Bathypelagic
Abyssopelagic
Shelfbreak
shallowwater assembly
deep diving, shelf & shelfbreakbreak assembly
surface feeding, oceanic assembly
25
0m
200m
1000m
2-3000m
Neritic OceanicEpipelagic
Mesopelagic
Bathypelagic
Cetaceans & seabirds utilise resources within the water column up to a depth of c. 3000m.
Oceanic seabirds, however, rarely dive particularly deep
27
Data visualization from MODIS on NASA’s AQUA spacecraft: a measure of global chlorophyll concentrations (data collected 2002-2004, 4-km resolution). Credit: NASA
© CJ Camphuysen Grey Phalarope
Grey Phalarope© H Verdaat
© CJ Camphuysen Sabine’s Gulls & White-capped Albatrosses
Cape Town, Feb 2001
Sooty shearwater tracks
From breeding grounds to rich foragingareas
Shaffer et al. 2006. Migratory shearwatersintegrate oceanic resources across the Pacific Ocean in an endless summer. PNAS 103: 12799-1280.
Shearwater megaflock in Unimak Pass, Alaska, Photo Kevin Bell 31
Seabird foraging techniques32
aerial
surface
subsurface
33
How to find food…..in an ‘empty’ environment with patchy resources?
• Visual cues detect prey or foraging flocks by eye
• Olfactory cues smell resources
• Auditory cues hear prey or produce/receive sound
• Habitat ‘understanding’ know situations that enhancefeeding opportunities
• Memory remember reliable locations
• Interpretation interpret other cues as likely feedinglocations
© CJ Camphuysen
35
• Visual cues go for any discontinuity in the greatemptiness of the ocean!
detection of any floating mattermonitor shippingvisit ripples and flotsam
• Visual cues
Scavenging at fishing vessel 37
• Visual cues
38
• Visual cues how to detect prey by eye
© CJ Camphuysen
Underwater prey is notoriously hard to detect and the field of vision of an overflying seabird is limited.Adverse weather will reduce the likelihood of preydetection.
39
• Visual cues go for conspecifics or competitors!
detect foraging flocks by eyemonitor the behaviour of conspecifics
off Schiermonnikoog, Aug 2013Harbour Porpoise (driver), Lesser Black-backed Gull, Northern Gannet, sandeels (prey) © H Verdaat
Natural feeding frenzy
40
Northern GannetNW Europe, Canada
• wintering • shelf break & shelf• widespread, abundant• c. 325,000 individuals• some fisheries
interactions• peak at low SST/SSS• natural feeding
frenzies• MSFA joiner
SSS SST
Depth
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.0W 17.5W 17.0W 16.5W 16.0W
Northern Gannet per km2
10.00 +
5.00 - 9.99
2.00 - 4.99
1.00 - 1.99
0.01 - 0.99
No birds
Blank = not surveyed
Density (birds/km2)
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.5W 18.0W 17.5W 17.0W 16.5W 16.0W
Northern Gannet / Fou de Bassan
26+
11-25
6-10
2-5
1
0
5
10
15
20
25
30
35
0
5
10
15
20
25
30
35
40
45
De
nsi
tie
s o
f b
ird
s (n
pe
r km
²)
Ob
serv
er
eff
ort
(km
²)
Effort
Gannet
0
5
10
15
20
25
30
35
40
45
50
0
10
20
30
40
50
60
70
De
nsi
tie
s o
f b
ird
s (n
pe
r km
²)
Ob
serv
er
eff
ort
(km
²)
Effort
Gannet
0
5
10
15
20
25
30
35
0-50 50-100 100-200 200-300 300-400 400-500 500-600 600-700 700-800 800-900 900-1000 >1000
Den
sity
(n k
m-2
)
gannets
1469 Northern Gannets were seen in association with fishing vessels. Substantial flocks:
07 Dec 2012 18°52.6’N, 16°38.2’W (372 individuals).07 Dec 2012 18°52.6’N, 16°37.5’W ( 37 individuals).07 Dec 2012 18°52.6’N, 16°37.1’W (480 individuals).07 Dec 2012 18°52.6’N, 16°37.1’W ( 40 individuals).07 Dec 2012 18°52.6’N, 16°36.8’W (368 individuals).
07 Dec 2012 18°52.6’N, 16°36.0’W ( 99 individuals).
13,417 Northern Gannets in natural feeding frenzies. Most substantial flocks (>100) :
08 Dec 2012, 20°07.5’N, 17°36.6’W (2570 individuals).08 Dec 2012, 20°07.5’N, 17°38.1’W (1700 individuals).04 Dec 2012, 19°30.1’N, 16°55.6’W (1397 individuals).04 Dec 2012, 19°30.1’N, 16°54.9’W (1200 individuals).04 Dec 2012, 19°30.1’N, 16°53.5’W ( 900 individuals).04 Dec 2012, 19°30.1’N, 16°54.2’W ( 770 individuals).08 Dec 2012, 20°07.5’N, 17°34.4’W ( 602 individuals).08 Dec 2012, 20°07.4’N, 17°22.4’W ( 500 individuals).08 Dec 2012, 19°52.5’N, 17°21.3’W ( 425 individuals).08 Dec 2012, 20°12.0’N, 17°39.2’W ( 407 individuals).04 Dec 2012, 19°37.4’N, 16°58.1’W ( 289 individuals).08 Dec 2012, 20°07.4’N, 17°40.4’W ( 142 individuals).08 Dec 2012, 19°54.6’N, 17°37.7’W ( 122 individuals).08 Dec 2012, 20°07.5’N, 17°33.7’W ( 120 individuals).04 Dec 2012, 19°30.1’N, 16°57.0’W ( 115 individuals).08 Dec 2012, 20°07.5’N, 17°33.7’W ( 115 individuals).08 Dec 2012, 19°52.5’N, 17°21.3’W ( 110 individuals).03 Dec 2012, 20°30.4’N, 17°26.4’W ( 106 individuals).
43
• Olfactory cues smell resources
Grubb T.C. 1972. Smell and foraging in shearwaters and petrels. Nature 237: 404-405.
European Storm-Petrel Hydrobates pelagicus Stormvogeltje
© CJ Camphuysen
Wilsons Storm-Petrel Oceanites oceanicus Wilsons Stormvogeltje
Camphuysen C.J. 2007. Where two oceans meet: offshore interactions of Great-winged Petrels Pterodromamacroptera and Leach's Storm petrels Oceanodroma leucorhoa off southern Africa. J. Ornithol. 148: 333-346.
Great-winged Petrel
Leach’s Storm-petrel
Camphuysen C.J. 2007. Where two oceans meet: offshore interactions of Great-winged Petrels Pterodromamacroptera and Leach's Storm petrels Oceanodroma leucorhoa off southern Africa. J. Ornithol. 148: 333-346.
Great-winged Petrel
Leach’s Storm-petrel
Relationships between the distribution of great winged petrels and Leach’s storm petrels and meso-scalehydrographic features in the Agulhas current retroflectionregion. The map shows sea surface height anomalies (m)
48
• Habitat ‘understanding’• Memory• Interpretation
49
• Habitat ‘understanding’• Memory• Interpretation
foto Bureau Waardenburg
50
• Habitat ‘understanding’• Memory• Interpretation
deep ocean (>1000m) shelf break (100-1000m) shelf sea (<100m)
Eastern boundary upwelling ecosystems
Coastal upwelling occurs in regions where winds push surface water off-shore and is replaced by deeper water.
NW African coast
The main coastal upwelling systems are found on the eastern side of the ocean basins
• Canary Current upwelling system
• Benguela Current upwelling system
• California Current upwelling system
• Peru Current upwelling system
Upwelling areas support a rich ecosystem
• Upwelled water is rich in nutrients like nitrogen and phosphorus.
• When upwelled water reaches the sun-lit zone, nutrients allow the growth of phytoplankton.
• Phytoplankton are grazed by zooplankton, small animals that are in turn eaten by fish
• Fish is eaten in turn by seabirds and marine mammals.
SST map across the Mauritania upwelling zone 15–17 July 2005.
The white dashed line shows the location of the shelf edge.
Source: Wynn &. Krastel 2012. Seabird 25: 47-53.
European Storm-petrelNW Europe, Mediterranean
• wintering, migrant • shelf sea, break• patchy distribution• c. 280,000 individuals• olfactory feeding,
communal?• low SST and SSS• at the periphery of
feeding frenzies
SSS SST
Depth
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.0W 17.5W 17.0W 16.5W 16.0W
European Storm Petrels per km2
5.00 +
3.00 - 4.99
2.00 - 2.99
1.00 - 1.99
0.01 - 0.99
No birds
Blank no counts
Density (n/km2)
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.5W 18.0W 17.5W 17.0W 16.5W 16.0W
European Storm-petrel / Océanite tempete
26+
11-25
6-10
2-5
1
0
1
2
3
4
5
6
7
8
0
5
10
15
20
25
30
35
40
45
<1
7.5
°C
17
.5-1
8.0
°C
18
.0-1
8.5
°C
18
.5-1
9.0
°C
19
.0-1
9.5
°C
19
.5-2
0.0
°C
20
.0-2
0.5
°C
20
.5-2
1.0
°C
>2
1.0
°C
Bir
ds
pe
r km
² (d
en
siti
es)
Ob
serv
er e
ffo
rt (k
m²)
Effort
Eur Storm-p
-2
0
2
4
6
8
10
12
0
10
20
30
40
50
60
70
35
.80
-36
.00
‰
36
.01
-36
.30
‰
36
.31
-36
.60
‰
36
.61
-36
.90
‰
36
.91
-37
.20
‰
37
.21
-37
.50
‰
37
.51
-37
.80
‰
37
.81
-38
.54
‰
Bir
ds
pe
r km
² (d
en
siti
es)
Ob
serv
er e
ffo
rt (k
m²)
Effort
Eur Storm-p
0
1
2
3
4
5
6
7
8
0-50 50-100 100-200 200-300 300-400 400-500 500-600 600-700 700-800 800-900 900-1000 >1000
Den
sity
(n k
m-2
)
European Storm-petrel
Band-rumped Storm-petrelMacaronesiaLeach’s Storm-petrelCanada, NW Europe
• wintering, migrant • oceanic• widespread, scarce• c. 66,000 individuals• high SST/SSS• periphery of feeding
frenzies• olfactory feeding• non-communal?
SSS SST
Depth
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.0W 17.5W 17.0W 16.5W 16.0W
Band-rumped Storm Petrels per km2
5.00 +
3.00 - 4.99
2.00 - 2.99
1.00 - 1.99
0.01 - 0.99
No birds
Blank no counts
Density (n/km2)
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.5W 18.0W 17.5W 17.0W 16.5W 16.0W
Band-rumped Storm-petrel / Océanite de Castro
26+
11-25
6-10
2-5
1
0
1
2
3
4
5
6
7
0
5
10
15
20
25
30
35
40
45
<1
7.5
°C
17
.5-1
8.0
°C
18
.0-1
8.5
°C
18
.5-1
9.0
°C
19
.0-1
9.5
°C
19
.5-2
0.0
°C
20
.0-2
0.5
°C
20
.5-2
1.0
°C
>2
1.0
°C
Bir
ds
pe
r km
² (d
en
siti
es)
Ob
serv
er e
ffo
rt (k
m²)
Effort
Band-r Stp
0
0.2
0.4
0.6
0.8
1
1.2
1.4
1.6
0
10
20
30
40
50
60
70
35
.80
-36
.00
‰
36
.01
-36
.30
‰
36
.31
-36
.60
‰
36
.61
-36
.90
‰
36
.91
-37
.20
‰
37
.21
-37
.50
‰
37
.51
-37
.80
‰
37
.81
-38
.54
‰
Bir
ds
pe
r km
² (d
en
siti
es)
Ob
serv
er e
ffo
rt (k
m²)
Effort
Band-r Stp
0
0.5
1
1.5
2
2.5
0-50 50-100 100-200 200-300 300-400 400-500 500-600 600-700 700-800 800-900 900-1000 >1000
De
nsi
ty (n
km
-2)
Band-rumped & Leach's Storm-petrel
H Verdaat
Grey PhalaropeHigh arctic
• wintering• shelf break• patchy distribution• c. 122,000 individuals• no fisheries
interactions• eddies/fronts (var SST)• at fronts and flotsam
concentrations• flock-feeder
SSS SST
Depth
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.0W 17.5W 17.0W 16.5W 16.0W
Grey Phalarope per km2
5.00 +
3.00 - 4.99
2.00 - 2.99
1.00 - 1.99
0.01 - 0.99
No birds
Blank no counts
Density (n/km2)
18.0N
18.5N
19.0N
19.5N
20.0N
20.5N
21.0N
18.5W 18.0W 17.5W 17.0W 16.5W 16.0W
Grey Phalarope / Phalarope à bec large
26+
11-25
6-10
2-5
1
-5
0
5
10
15
20
25
30
0
5
10
15
20
25
30
35
40
45
<1
7.5
°C
17
.5-1
8.0
°C
18
.0-1
8.5
°C
18
.5-1
9.0
°C
19
.0-1
9.5
°C
19
.5-2
0.0
°C
20
.0-2
0.5
°C
20
.5-2
1.0
°C
>2
1.0
°C
Bir
ds
pe
r km
² (d
en
siti
es)
Ob
serv
er e
ffo
rt (k
m²)
Effort
Phalaropes
-2
0
2
4
6
8
10
12
14
16
18
0
10
20
30
40
50
60
70
35
.80
-36
.00
‰
36
.01
-36
.30
‰
36
.31
-36
.60
‰
36
.61
-36
.90
‰
36
.91
-37
.20
‰
37
.21
-37
.50
‰
37
.51
-37
.80
‰
37
.81
-38
.54
‰
Bir
ds
pe
r km
² (d
en
siti
es)
Ob
serv
er e
ffo
rt (k
m²)
Effort
Phalaropes
0
5
10
15
20
25
30
0-50 50-100 100-200 200-300 300-400 400-500 500-600 600-700 700-800 800-900 900-1000 >1000
Den
sity
(n k
m-2
)
phalaropes
Observer effort (km²) Grey Phalaropes (n km-2) Band-rumped Petrels (n km-2) Europ Storm Petrels (n km-2)
°C\‰ 35 36 37 38 °C\‰ 35 36 37 38 °C\‰ 35 36 37 38 °C\‰ 35 36 37 38
16 3.9 0.4 16 0.0 0.0 16 0.0 0.0 16 0.0 0.0
17 0.8 50.4 17 0.0 0.1 17 0.0 0.0 17 0.0 5.0
18 1.6 40.4 16.0 1.1 18 0.0 1.3 11.2 0.0 18 0.0 0.2 0.4 0.0 18 0.6 1.7 0.1 0.0
19 13.2 35.4 19 9.6 7.0 19 0.0 0.2 19 0.2 0.9
20 21.3 20 14.7 20 1.1 20 2.0
21 1.1 21 0.0 21 6.1 21 0.9
H Verdaat
58
• Habitat ‘understanding’• Memory• Interpretation
59© CJ Camphuysen
Foraging marine mammals &seabird – cetacean interactions
© L Meeuwisse
60
Surface feeding seabirds take advantage of fish driven to the surface by underwater predators
Herding
Most attacked prey fish willpromptly form a highly compact mass of animals,
a fish ball
Fish balls tend to intimidate predators and provide relative safetyfor individual fish
61
Herding
http://i.telegraph.co.uk/telegraph/multimedia/archive/01500/bait-ball-dolphins_1500165i.jpg
Effective predators drive fish towards a physical barrier (e.g. the sea surface)
62
63
64
65
66
68
Foraging tactics and multi-species feeding frenzies
69
from: Camphuysen & Webb 1999
Surface feeding seabirds take advantage
of fish driven to the surface by underwater predators
70
Camphuysen C.J. & A. Webb 1999. Multi-species feeding associations in North Sea seabirds: jointly exploitinga patchy environment. Ardea 87: 177-198.
71
Surface feeding seabirds take advantage of fish driven to the surface by underwater predators
72
Surface feeding seabirds take advantage of fish driven to the surface by underwater predators
73
Herding
http://i.telegraph.co.uk/telegraph/multimedia/archive/01500/bait-ball-dolphins_1500165i.jpg
Nottestad L. & B.E. Axelsen 1999. Herring schooling manoeuvres in response to killer whale attacks. Can. J. Zool. 77: 1540-1546.
The antipredator behaviour of herring was investigated during repeated attacks by killer whales in Tysfjord in northwestern Norway using a high-resolution (455 kHz) multibeam sonar.
Ten different types of predator-prey interactions were recorded.
Antipredator responses included "split," "hourglass," "vacuole," "bend," "dive," "herd," and "fountain."
Large attacked schools demonstrated a different repertoire of antipredatormanoeuvres than small ones and were less likely to be attacked.
Herring schools with a cross section exceeding 460 m² were not attacked by killer whales.
Attacked schools were significantly more circular (p < 0.0001) and had higher relative densities (p < 0.05) than schools that were not attacked.
74
http://i.telegraph.co.uk/telegraph/multimedia/archive/01500/bait-ball-dolphins_1500165i.jpg
Nottestad L. & B.E. Axelsen 1999. Herring schooling manoeuvresin response to killer whale attacks. Can. J. Zool. 77: 1540-1546.
75
Herding
Fish herding predators usually act in groups in close co-operation: simultaneous diving, concerted actionunder water
Herding
© CJ Camphuysen
76
Does being “baldly patterned” help….?Herding
Ryan P.G., Wilson R.P. & Cooper J. 1987. Intraspecific mimicry and status signals in
juvenile Africian penguins. Behav. Ecol. Sociobiol. 20: 69-76.
Herding
poor fish-herding properties excellent fish herding properties
80© CJ Camphuysen
Trawling or Herding ?
Humpback Whale
Brydes Whale
Brydes Whale
© CJ Camphuysen
83
Humpback Whale
Spy-hopping whales have been found to actively search for seabird feeding frenzies and join the feast
84
85
15
The marine food web
The concept of central-place foraging
86
Simple rule of thumb:An animal should gain energy during a foraging trip,energy for maintenance, growth, and/or reproduction
The costs of travel (locomotion), handling, digestion and prey carrying should be outweighed by the food intake
Energy requirements: FMR + the needs for partner & offspring during breeding
87
Satellite tracks from 184 Northern Gannets show that foraging birds direct their movements away from neighboring colonies (colors correspond to different colonies).
Wakefield et al. 2013. Space partitioning without territoriality in Gannets. Science 341: 68-70.
The concept of central-place foraging
The information centre hypothesis suggests that birds that breed in colonies may exchange information about the location of (ephemeral) food patches
Antarctic Prions © CJ Camphuysen
Ward & Zahavi 1973. The importance of certain assemblages of birds as "information-centres" for food-finding. Ibis 115: 517-534. 88
The concept of central-place foraging
Climate change and shifts in resources
Extreme temperatures in the North Pacific are cancelling out El Niño and wreaking their own climate havoc
New Scientist October 2014
90
Anker-Nilssen T. 1987. The breeding performance of Puffins Fratercula arctica on Rost, northern Norway in 1979-1985. Fauna Norv. Ser. C., Cinclus10: 21-38.
Climate change and shifts in resources
Seabirds around Iceland have been declining in the last 10-15 years:Brünnich’s guillemot by 44% and there are 30% fewer guillemots and fulmars.
The main reason: changes in ocean currents which have led to a crash in sandeels
Climate change and shifts in resources
Norway was one of the countries that caught the highest numbers of bluefin tuna in Europe in the 1950s and 1960s, but the fishery was over by the mid-1980s.
In 2008, Norway issued on a ban on the fishery.
Climate change and shifts in resources
Climate change and shifts in resources
The world’s oceans: Never short of surprises
Atlantic Grey Whale Eschrichtius robustus
Extinct
Marine Sciences I (2013) Kees (CJ) Camphuysen Royal NIOZ Texel Marine Mammals 94
Marine biologists are scratching their heads this week, trying to figure out how a grey whale, normally found only in the Pacific Ocean, has been spotted off the coast of Israel, 10 May 2010.
The solitary whale, measuring some 12 metres (39 feet) and weighing around 20 tonnes, was first sighted off Herzliya on the coast of the Mediterranean, on Saturday.
A mysterious gray whale sighted off the coast of Israel in the Mediterranean Sea has been seen again off the north east coast of Spain.
The second sighting, made 23 days and 3000km after the first, has continued to perplex whale experts.
http://news.bbc.co.uk/earth/hi/earth_news/newsid_8672000/8672970.stm
http://news.bbc.co.uk/earth/hi/earth_news/newsid_8729000/8729064.stmMarine Sciences I (2013) Kees (CJ) Camphuysen Royal NIOZ Texel Marine Mammals 95
Hypothetical route
Atlantic Grey Whale overlooked odd 400 years?
Pacific Grey Whale of a lengthy journey?
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97
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