influence of hypoxia on the distribution, behavior, and foraging of zooplankton and planktivorous...
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Influence of hypoxia on the distribution, behavior, and foraging of zooplankton and planktivorous fish in centralLake Erie: Field observations & future directions
Hank Vanderploeg, GLERLStuart Ludsin, GLERLSteve Pothoven, GLERLTomas Höök, CILER Univ. of MichiganJames Roberts, Univ. of MichiganSteve Ruberg, GLERLJoann Cavaletto, GLERLJames Liebig, GLERLGregory Lang, GLERLStephen Brandt, GLERL
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Hypoxia is an old problem in freshwater—Results forCyclops bicuspidatus (Einsle 1965)
This species is very tolerant of low oxygen (~ 0.1mg/L)
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1. Hypoxia will disrupt vertical migration behavior
– Reduce time spent on bottom
2. Hypoxia will influence horizontal movement
– Fish will move into oxygenated, shallow nearshore zones
3. Hypoxia will reduce availability of prey, both ZP & benthic macroinvertebrate prey
– ZP use hypoxia as a refuge from predation– Hypoxia reduces benthic prey abundance
4. Fish consumption & condition will decline
Original Lake ErieFish-Centric Hypotheses
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Playing chess with death—a zooplankton-centric view
Scene from Bergman’s “The Seventh Seal”
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Death normally comes in two forms: predation and starvation
• Zooplankton vertical migration is strategy to minimize overlap with visually preying invertebrate and vertebrate (fish) predators—conspicuous or unprotected (spineless) zooplankton move to lower light levels
• Move into upper favorable (temperature and food) areas at night.
• Predator abundance is assessed by kairomones.• When many predators, the zooplankter (prey)
must play chess to avoid overlap.
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The Great Lakes have both visual invertebrate & and vertebrate predators—Lake Michigan example
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Playing chess with death—the piscine players
Scene from Bergman’s “The Seventh Seal”
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Rainbow Smelt August 2005
0.00
0.25
0.50
0.75
1.00
W' Night
Day
Emerald Shiner August 2005
0.00
0.25
0.50
0.75
1.00
W' Night
Day
USGS-NAS
Emerald shiner:Epilimnetic planktivore
Rainbow Smelt:Planktivore-benthivore
Dominant planktivores of Lake Erie and their Vanderploeg & Scavia (1979) selectivity coefficients (W´) pre-hypoxia
Prey size
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Hypoxia, another form of death, alters the game—some hypotheses:
• Differential tolerance of zooplankton to hypoxia allows some species to enter the hypoxic zone to escape predators—the refuge
• Others will be forced out and trapped in lighted areas above—the hypoxia-light trap.
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-83.5 -82.5 -81.5 -80.5 -79.5
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-83.5 -82.5 -81.5 -80.5 -79.5
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DissolvedOxygen(mg/l)
0
3
6
9
12
September
Diel Station B
August
Lake Erie
Some results before and after major hypoxia will give us some insights
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General Methods—What we did
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• Trawling (fish species & samples for diet & ration work)
• Zooplankton net and pump sampling (zooplankton)
• Ponar sampling (benthic macroinvertebrates)
• Zooplankton• Temperature• Dissolved oxygen• Light levels• Chlorophyll a
FishBiomass
Introduction to Study Systems & General Methods
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Lake Erie Field Program (IFYLE 2005)
Diel (24-hr)Transect (day-night)
Source: Don Coles
EPA-GLNPOR/V Lake Guardian (180’)
NOAA-GLERLR/V Laurentian (80’)
Transect BDiel Station B
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0 5 10 15 20 25D
epth
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)0
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Water Column Pumping Method
1 min. ea. depth
1 min. ea. depth
5 min.ea.
DO (mg/L)
Water Temp (oC)
2 min. ea. depth
Sept. 2005
shooting for pumping 1 cubic meter of water
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DissolvedOxygen(mg/l)
0
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12
-83.5 -82.5 -81.5 -80.5 -79.5
41.5
42
42.5
43-83.5 -82.5 -81.5 -80.5 -79.5
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-83.5 -82.5 -81.5 -80.5 -79.5
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Transect B
Ho 2: Hypoxia will alter horizontal distribution of abundance– Fish will move into oxygenated, shallow nearshore zones
September
August
October
Lake Erie
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41.7 41.8 41.9 42 42.1
20
10
0
0
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12
- 1 1 0
- 9 0
- 7 0
- 5 0
- 3 0
Latitude (degrees)
Dep
th (
m)
Day
41.7 41.8 41.9 42 42.1
20
10
0
NightTemp(º C)
DO(mg/l)
Fish(dB)
6
1 4
2 2
3 0
(August – Pre-Hypoxia)
41.7 41.8 41.9 42 42.1
20
10
0
41.7 41.8 41.9 42 42.1
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0
41.7 41.8 41.9 42 42.1
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20
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0
Ho 2: Hypoxia will alter horizontal distribution of abundance
Lake Erie
Ludsin, Vanderploeg & Ruberg, unpub
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41.7 41.8 41.9 42 42.1
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- 1 1 0
- 9 0
- 7 0
- 5 0
- 3 0
Latitude (degrees)
Dep
th (
m)
Temp(º C)
DO(mg/l)
Fish(dB)
6
1 4
2 2
3 0
41.7 41.8 41.9 42 42.1
20
10
0
41.7 41.8 41.9 42 42.1
20
10
0
41.7 41.8 41.9 42 42.1
20
10
0
(September – Peak Hypoxia)
Ho 2: Hypoxia will alter horizontal distribution of abundance
Day
41.7 41.8 41.9 42 42.1
20
10
0
Night
Lake Erie
Ludsin, Vanderploeg & Ruberg, unpub
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0
3
6
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- 1 1 0
- 9 0
- 7 0
- 5 0
- 3 0
Latitude (degrees)
Dep
th (
m)
Temp(º C)
DO(mg/l)
Fish(dB)
6
1 4
2 2
3 0
41.6 41.7 41.8 41.9
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0
41.6 41.7 41.8 41.9
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0
41.7 41.8 41.9 42 42.1
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10
0
41.7 41.8 41.9 42 42.1
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10
0
41.7 41.8 41.9 42 42.1
20
10
0
(October – Post Hypoxia)
Ho 2: Hypoxia will alter horizontal distribution of abundance
Day
41.7 41.8 41.9 42 42.1
20
10
0Night
– Reject: Fish move into oxygenated waters, but offshore
Lake Erie
Ludsin, Vanderploeg & Ruberg, unpub
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Playing chess with death—Insights from pre-hypoxia (control) & hypoxia distributions and prey selection
Scene from Bergman’s “The Seventh Seal”
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Diel B, Aug 17, 01:00 EDT
0
5
10
15
20
25
0 5 10 15 20 25
Chl, DO, Zoop, Temp
De
pth
(m
)
0 100 200 300 400 500PAR
Fish biomass(relative)
Zoomass (10 ug/L)
Chl (ug/L)
DO (mg/L)
Temp ('C)
PAR (uE/m2/s)
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Diel B, Aug 17, 13:00 EDT
0
5
10
15
20
25
0 5 10 15 20 25Chl, DO, Zoop, Temp
De
pth
(m
)
0 200 400 600 800 1000 1200PAR
Fish biomass(relative)
Zoomass (10 ug/L)
Chl (ug/L)
DO (mg/L)
Temp ('C)
PAR (uE/m2/s)
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Copepods mg . m-3
0 10 20 30 40
dep
th
0
4
8
12
16
20
24
Cladocerans mg . m-3
0 100 200
Lake Erie B 8-17-05 DIEL 02:00
BosminaEubosminaDaphnia mendotaeD. longiremis
Predatory Cladocerans mg . m-3
0 10 20 30
4.8 mg/L DO
EPI
META
HYPO
4.8 mg/L DO 4.8 mg/L DO
LeptodoraBythotrephesCercopagis
DiacyclopsMesocyclopsTropocyclopsDiaptomidsEpischuranauplii
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Copepods mg . m-3
0 50 100
dep
th
0
4
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12
16
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24
DiacyclopsMesocyclopsTropocyclopsDiaptomidsEpischuranauplii
Cladocerans mg . m-3
0 20 40
Lake Erie B 8-17-05 DIEL 14:00
BosminaEubosminaDaphnia mendotaeD. longiremisD. retrocurva
Predatory Cladocerans mg . m-3
0 1 2
EPI
META
HYPO
4.8 mg/L DO 4.8 mg/L DO 4.8 mg/L DO
LeptodoraBythotrephes
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Diel B, Sept 18, 03:00 EDT
0
5
10
15
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25
0 5 10 15 20 25
Chl, DO, Zooplankton, Temp
De
pth
(m
)
0 100 200 300 400 500PAR
Fish biomass(relative)
Zoomass (10 ug/L)
Chl (ug/L)
DO (mg/L)
Temp ('C)
PAR (uE/m2/s)
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Diel B, Sept 17, 15:00 EDT
0
5
10
15
20
25
0 5 10 15 20 25
Chl, DO, Zooplankton, Temp
De
pth
(m
)
0 100 200 300 400 500PAR
Fish biomass(relative)
Zomass (10 ug/L)
Chl (ug/L)
DO (mg/L)
Temp ('C)
PAR (uE/m2/s)
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Predatory Cladocerans mg . m-3
0 1 2 3Copepods mg . m-3
0 100 200
dep
th
0
4
8
12
16
20
24
Cladocerns mg . m-3
0 20 40 60 80
Lake Erie B 9-18-05 DIEL 02:00
DiacyclopsMesocyclopsTropocyclopsDiaptomidsEpischuranauplii
BosminaEubosminaDaphnia mendotaeD. longiremisD. retrocurvaDiaphanasoma
Leptodora
upper epi
lower epi
meta
hypo
1.2 mg/L DO 1.2 mg/L DO 1.2 mg/L DO
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Copepods mg . m-3
0 20 40 60 80
dep
th
0
4
8
12
16
20
24
Cladocerans mg . m-3
0 20 40
Lake Erie B 9-17-05 DIEL 14:00
DiacyclopsMesocyclopsTropocyclopsDiaptomidsEpischuranauplii
Predatory Cladocerans mg . m-3
0.0 0.5 1.0
1.2 mg/L DO 1.2 mg/L DO 1.2 mg/L DO
upper epi
lower epi
meta
hypo
BosminaEubosminaDaphnia mendotaeD. longiremisD. retrocurvaDiaphanasoma
Leptodora
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Rainbow Smelt August 2005
0.00
0.25
0.50
0.75
1.00
W' Night
Day
Emerald Shiner August 2005
0.00
0.25
0.50
0.75
1.00
W' Night
Day
USGS-NAS
Emerald shiner:Epilimnetic planktivore
Rainbow Smelt:Planktivore-benthivore
Selectivity coefficient of Vanderploeg & Scavia (W´) for Emerald shiner and Rainbow Smelt in August 2005
Prey size
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Rainbow Smelt September 2005
0.00
0.25
0.50
0.75
1.00
W' Night
Day
Emerald Shiner September 2005
0.00
0.25
0.50
0.75
1.00
W' Night
Day
USGS-NAS
Emerald shiner:Epilimnetic planktivore
Rainbow smelt:Planktivore-benthivore
Prey size
Selectivity coefficient of Vanderploeg & Scavia (W´) for emerald shiner and rainbow smelt in September 2005
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What’s going on down there?
Present status:Heavy emphasis in IFYLE Hypoxia study on upper
food web (fish and location of fish food)We do know, however:• Mesozooplankton and microzooplankton
distribution relative to hypoxia response is species specific
• Microzooplankton grazing dominates during the summer
• Bacteria-based food web becomes important in hypoxic zone
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What’s going on down there?
For the development of a conceptual framework we’d like to know:
• What is the minimum oxygen concentration a zooplankter (species by species) is willing to enter yet survive under various predation risk scenarios?
• How does feeding and behavior vary with oxygen concentration?
• What is the joint distribution of meso-and microzooplankton around hypoxic zones
• How is production and predation risk affected?
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We know something about Daphnia foraging in hypoxic areas but nothing for copepods, the dominants in the Great Lakes, or for visual
invertebrate predators
From Heisey & Porter (1977)
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Some possible lab approaches to define spatial
rules of food web assembly (“indirect effects”)
• Observe location of position of zooplankton in laboratory water columns with gradients of light, temperature, kairomones of potential predators & oxygen
• Directly observe behavior and foraging in hypoxic water columns.
• Observe effect of hypoxia on visual predation (both invertebrate & vertebrate)—have predators watch TV
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Inside the lab
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Outside the lab: keeping the predator in focus