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Role of zooplankton in marine ecosystemsand modelling perspectivesg p p
Tom AndersonNational Oceanography Centre, Southampton, UK
ASP Researcher Workshop NCARASP Researcher Workshop, NCARAug 6-10th 2013
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Role of zooplankton in marine ecosystemsp yand modelling perspectives
Introduction: bottom up (BU) vs top down (TD) control of marine ecosystems
Zooplankton control of phytoplankton blooms Zooplankton control of phytoplankton blooms
Zooplankton control of plankton community structure
Modelling zooplankton: state-of-the-art
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Role of zooplankton in marine ecosystemsRole of zooplankton in marine ecosystemsand modelling perspectives
Introduction: bottom up (BU) vs top down (TD) control of marine ecosystems
Zooplankton control of phytoplankton blooms Zooplankton control of phytoplankton blooms
Zooplankton control of plankton community structure
Modelling zooplankton: state-of-the-art
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evapotranspiration
NPPtree PFT2
fluxesCH2O
NPPtree PFT1
grasses
NPP
soilrespirationdecomposition
precipitation
soil
pp
uptakeuptake
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Sea surface temperature Chlorophyllp p y
http://earthobservatory.nasa.gov
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16 most abundant organized by “functional group”
Prochlorococcus Prochlorococcus analogsanalogs
SynechococcusSynechococcus & small eukaryotes& small eukaryotes
DiatomsDiatoms.
Other large eukaryotesOther large eukaryotes
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Plant animal warsPlant-animal wars
spinesspinescoloniestoxinstoxins
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Daksalov (2002): “Overfishing drives a trophic cascade in the Black sea”( ) g p
Change in dominance across the pelagic food chain since the depletion oftop-predators after 1970: model results
trophic level before 1970 after 1970
top predators
planktivores
resource limited
consumer controlled
-
resource limitedp
zooplankton
phytoplankton
resource limited
consumer controlled
consumer controlled
resource limitedphytoplankton consumer controlled resource limited
Mar. Ecol. Prog. Ser. 225, 53-63
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Green world hypothesis
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The “Biological Pump”The “Biological Pump”The Biological PumpThe Biological Pump
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Export
No simple relationship with primary production
Much export occurs during bloom events
Is related to ecosystem structure
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Active flux
Bianchi et al. (2013)
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Detritus turnover
A d d T (2010)Anderson and Tang (2010)
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Role of zooplankton in marine ecosystemsRole of zooplankton in marine ecosystemsand modelling perspectives
Introduction: bottom up (BU) vs top down (TD) control of marine ecosystems
Zooplankton control of phytoplankton blooms Zooplankton control of phytoplankton blooms
Zooplankton control of plankton community structure
Modelling zooplankton: state-of-the-art
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Plankton stocks are a balance of opposing forces :accumulation and degradation, consumption and regenerationaccumulation and degradation, consumption and regeneration
(Riley, 1941)
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Pgrowth grazing Gordon Riley
Ph (1 k z )L LpI0
= (Ph – R – G)PdPdt
Ph = (1-e-k.zeu)LNLVpI0kzeu
TR = R0erT
photosynthesis grazingG = gZ
Riley (1946)
respiration
Riley (1946)
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Georges Bank simulationg
P ZP, Z
monthmonth
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maximum growth and grazing rates
MEDUSA
0.5 d-1
2.0 d-1
1 7 d 11.7 d-11.6 d-1
Yool et al. (2010, 2013)
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Annual cycle of mixed layer depthAnnual cycle of mixed layer depth
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India 60N 20W
KERFIX 50S 68E
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India 60N 20W
KERFIX 50S 68E
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60N 20W planktondiatoms mesozooplankton
60N 20W planktonnon-diatoms microzooplankton
plankton biomassl N
m-3
mm
ol
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growth60N 20W plankton
g
grazingd-1
non-diatoms diatoms
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60N 20W planktondiatoms mesozooplankton
no grazing planktonnon-diatoms microzooplankton
plankton biomassl N
m-3
mm
ol
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India 60N 20W
KERFIX 50S 68E
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Kerfix planktondiatoms mesozooplankton
Kerfix planktonnon-diatoms microzooplankton
plankton biomassl N
m-3
mm
ol
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growth
Kerfix planktong
grazingd-1
non-diatoms diatoms
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Kerfix planktondiatoms mesozooplankton
Kerfix,no grazing
planktonnon-diatoms microzooplankton
plankton biomassl N
m-3
mm
ol
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grazingplankton
no grazing
grazing
non diatoms it tnon-diatoms nitrate
ol N
m-3
mm
o
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Role of zooplankton in marine ecosystemsRole of zooplankton in marine ecosystemsand modelling perspectives
Introduction: bottom up (BU) vs top down (TD) control of marine ecosystems
Zooplankton control of phytoplankton blooms Zooplankton control of phytoplankton blooms
Zooplankton control of plankton community structure
Modelling zooplankton: state-of-the-art
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PlankTOM5
li h hmixed
coccolithophores diatomsphytoplankton
dissolveddissolvedorganic matter PO4 Fe SiO3
microzooplankton mesozooplanktonsmalldetritus
largedetritus
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1 0
0.8
1.0 BS
IvMM
ntak
e, d
-1
0 4
0.6MM
in
0.2
0.4
0 1 2 3 40.0
prey densityprey density
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diatoms mg chl m-3diatoms, mg chl m
March MayMM March-May B
0 60.6
0
S IV
(Anderson at al., 2010)
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Rosie Fisher (yesterday):
“The juxtaposition of numerous complex models createsTh i l f h l i l b h i ”The potential for pathological behaviour”
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coccolithophores mg chl m-3coccolithophores, mg chl m
March MayMM March-May B
0.4
0
S IV
(Anderson at al., 2010)
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microzooplankton mmol C m-3microzooplankton, mmol C m
March MayMM March-May B
0.8
0
S IV
(Anderson at al., 2010)
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mesozooplankton mmol C m-3mesozooplankton, mmol C m
March MayMM March-May B
5
0
S IV
(Anderson at al., 2010)
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Primary production g C m-2 d-1Primary production, g C m d
March MayMM March-May B
0.5
0
S IV
(Anderson at al., 2010)
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export g C m -2 d-1export g C m d
March MayMM March-May B
0.2
0
S IV
(Anderson at al., 2010)
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Role of zooplankton in marine ecosystemsRole of zooplankton in marine ecosystemsand modelling perspectives
Introduction: bottom up (BU) vs top down (TD) control of marine ecosystems
Zooplankton control of phytoplankton blooms Zooplankton control of phytoplankton blooms
Zooplankton control of plankton community structure
Modelling zooplankton: state-of-the-art
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Trophic transferingested
substrates
assimilationcosts
pelletsassimilation CO2
assimilated digestionsubstrates
moultmaintenance CO2 PO4
basal metabolism
maintenance
remainingsubstrates
protein turnover
t f
newbiomassproduction CO2
release
costs ofgrowth
growth (incl. reproduction)
excess C,P CO2 orDOC
PO4 orDOP
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Nit C b
pelletsCO2,NH4
Anderson & Hessen 95 ERSEM Anderson & Hessen 95 ERSEM
Nitrogen Carbon growth
1
0
frac
tion
Pahlow 03 Pahlow 03
0
1
n
0
frac
tion
Anderson et al. 05 HadOCC Anderson et al. 05 HadOCC1
on
food C/N4 20 food C/N4 20 food C/N4 20 food C/N4 200
frac
tio
food C/N4 20 food C/N4 20 food C/N4 20 food C/N4 20
Anderson et al. (in press)
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Anderson et al. (in press)
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1.0 B
-1
0.8
BS
IvMM
inta
ke, d
-
0.4
0.6i
0.2
0 1 2 3 40.0
prey density
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Loss terms and closure
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Complex models
ocean surface
Complex models
zooplanktongrazing
phytoplankton
ocean surface
euphotic zone NPP
microzooplankton mesozooplankton
g g
defecation
coccolithophores diatomsmixed
phytoplankton
“messy isinking
nutrients non-living organic matter
excretiondefecation,mortalityuptake
remineralisation
messyfeeding” grazing
mortality
sinking
PO4 Fe SiO3dissolved
organic mattersmall
detrituslarge
detritus
deep ocean mixing mixing sinking
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Poorly understood ecology
ingestedsubstrates
assimilation
pelletsassimilation CO2
costsassimilatedsubstrates
digestion
basal metabolism
moultmaintenance CO2
remaining
PO4
protein turnovergsubstrates
p
newbiomassproduction CO2
costs ofgrowth
biomassproduction
growth (incl reproduction)release
excess C,P CO2 orDOC
PO4 orDOP
growth (incl. reproduction)
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Aggregation
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Simplification
Calanus finmarchicus life history
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All in the interactions
coccolithophores mg chl m-3coccolithophores, mg chl m
MM March-May B
S IVV
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Lack of validation
Insufficient data
Too many free parameters
“What little validation is undertaken will always tend to overstate the case for belief in the model results”
(Wainwright, 2003)
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Conclusions
BU and TD combine to determine ecosystem structure andassociated fluxes such as export
The key is to understand the coupling of phytoplankton andl k di d b h i l izooplankton as mediated by physical environment
M d l k bl i i h i i f Models are remarkably sensitive to the parameterisation of zooplankton
The parameterisation of zooplankton in models is fraught withdifficulty, especially in complex models