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National Ecosystem Modeling Workshop (NEMoW) August 29-31 2007, NMFS Santa Cruz. Review of major ecosystem model classes. Éva Plagányi Dept. of Maths & Applied Maths, University of Cape Town. With thanks to Doug Butterworth and MARAM. - PowerPoint PPT Presentation
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Review of major ecosystem model classes
Éva PlagányiDept. of Maths & Applied Maths,
University of Cape TownReference: Plagányi 2007. Models for an Ecosystem Approach to Fisheries. FAO Fisheries
Technical paper 477
Report of Modelling Ecosystem Interactions for Informing an Ecosystem Approach to Fisheries: Best Practices in Ecosystem Modeling, Tivoli, July 3-6, 2007
National Ecosystem Modeling Workshop (NEMoW)August 29-31 2007, NMFS Santa Cruz
With thanks to Doug Butterworth and MARAM
POKOK BAHASAN
Ecosystem Model Objectives Ecosystem Model Types Questions for EAF* Modelling Ecosystem Model Classification Considerations in Model Building and the
Best Practice Approach Role of Management Procedures Data requirements Conclusions
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem Models and Management Advice1. Conceptual/understanding: of the structure, functioning and
interactions of the ecosystem, or sub-system, under consideration. May not be used explicitly in decision-making or scientific advice but forms the underlying context for any detailed management planning and decision-making
2. Strategic decisions: linked to policy goals and are generally long-range, broadly-based and inherently adaptable
3. Tactical decisions: aimed at the short-term (e.g. next 3-5 years), linked to an operational objective and in the form of a rigid set of instructions e.g. tactical decision to change quota
Ecosystem models generally intended to complement not replace single-species assessment models
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Tipe-tipe Model
I. Whole ecosystem models: models that attempt to take into account all trophic levels in the ecosystem
II. Minimum Realistic Models (MRM): limited number of species most likely to have important interactions with a target species of interest
III. Dynamic System Models (Biophysical): represent both bottom-up (physical) and top-down (biological) forces interacting in an ecosystem
IV. Extensions of single-species assessment models (ESAM): expand on current single-species assessment models taking only a few additional inter-specific interactions into account
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem ModelsI. Whole ecosystem models
MODEL NAMEEwE and ECOSPACE
Ecopath with Ecosim
ATLANTIS ATLANTISIGBEM Integrated
Generic Bay Ecosystem Model
INVITRO INVITROGEEM General
Equilibrium Ecosystem Model
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem Models- plankton focus (NPZ-fish)
MODEL NAMEERSEM II European
Regional Seas Ecosystem Model
SSEM Shallow Seas Ecological Model
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem ModelsII. Minimum Realistic Models
MODEL NAMEMRM Minimally Realistic
Models
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem ModelsII. Minimum Realistic Models
MODEL NAMEGADGET Globally applicable
Area Disaggregated General Ecosystem Toolbox
BORMICON BOReal Migration and CONsumption model
MULTSPEC Multi-species model for the Barents SeaSimplified version is AGGMULT which is also connected to a ECONMULT - a model describing the economies of the fishing fleet
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem ModelsII. Minimum Realistic Models cont.
MODEL NAMEMSVPA and MSFOR(and derivatives)
Multi-species Virtual Population Analysis; Multi-species Forecasting Model
MSM Multi-species Statistical Model
IBM Individual-Based Models
Bioenergetic/allometric
e.g. Koen-Alonso & Yodzis 2005SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem Models- Antarctic Models
MODEL NAMEFOOSA Previously KPFM
(Krill- Predator-Fishery Model)
SMOM Spatial Multi-species Operating Model
EPOC Ecosystem Productivity Ocean Climate model
Other CCAMLR models e.g. Mori & Butterworth 2005, 2006
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem ModelsIII. Dynamic System Models
MODEL NAMESEAPODYM Spatial Ecosystem
and Population Dynamics Model
OSMOSE Object-oriented Simulator of Marine ecOSystem Exploitation
SystMod System Model for the Norwegian and Barents Sea
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ecosystem ModelsIV. Extended Single-Species Models
MODEL NAMEESAM Extended Single-Species
Models e.g. Livingston and Methot 1998; Hollowed et al. 2000; Tjelmeland and Lindstrøm 2005
SEASTAR Stock Estimation with Adjustable Survey observation model and TAg-Return data
Target Species
Predator
Catch Catch
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Questions for EAF Modelling Issues pertaining to the management of target and related
species:
• Impact of a target species on other species in the ecosystem?
• Limitations of single-species-based assessment • Targeting of relatively unexploited species • What are the impacts of retained by-catch?• What is the effect on top predators of removing
the predators themselves and their prey? • What is the extent of competition between
fisheries and species of concern such as marine mammals, turtles, seabirds and sharks.
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Questions for EAF Modelling
Issues pertaining to species:
• What are the impacts of fishing on biodiversity?
• What are the impacts of commencing fishing on a previously unexploited species about which little is known.
• Effects of the introduction of non-native species.
• What are the impacts of non-retained by-catch?
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Questions for EAF Modelling Environmental and unintentional impacts on ecosystems
• Effects of physical/environmental factors on the resources on which fisheries depend.
• Changes in ecosystem state, e.g. regime shift, change to a less productive/less desirable state.
• Anthropogenic effects.• Effects of habitat modification e.g. trawling
damaging benthic habitats
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
1 2 3 4 5 6 7 8 9 10RESEARCH QUESTION/
MODEL
Ecopath with Ecosim and ECOSPACE IGBEM ATLANTIS INVITRO ERSEM II SSEM KPFM*
MRM e.g. Punt and Butterworth (1995)
MSVPA and MSFOR MSM
1a. Understanding - subset of ecosystem1b. Understanding - complete ecosystem
2. Impact of target species
3. Effect of top predators4. Competition: marine mammals - fisheries
5. Rebuilding depleted fish stocks6. Biases in single-species assessment7. Ways to distribute fishing effort among fisheries
8. Under-exploited species
9. Change in ecosystem state10. Spatial concentration of fishing11. Environmental/physical effects12. Effects of habitat modification
13. Effects of by-catch14. Introduction of non-native species* Still being developed
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Pg. 4
NO E
NVIR
ONM
ENT
ENVIRONMENT
AGE STRUCTUREAGE STRUCTURE
Biological interactions described
Predator prey feedback
Handles the environment and lower trophic levels
Technical interaction models MSYPRMurawski 1984
Predators added to single-species models e.g. SEASTARGulland 1983; Livingston and Methot 1998; Hollowed et al. 2000; Plagányi 2004; Tjelmeland and Lindstrøm 2005
No
No
Yes
Yes
Handles age/size structureHandles age
structure
No Yes
Aggregate system models e.g. EwE, SKEBUB, SSEM
Spatial dynamic systems models e.g. ATLANTIS, ERSEM, SEAPODYM
Multispecies Production Models e.g. Horbowy 2005
YesNo
Dynamic multi-species models BORMICON, GADGET, MRMs, MSVPA& MSFOR, MSM, MULTSPEC, OSMOSE
No
Handles spatial structure
Yes
No
Dynamic systems models e.g. some recent EwE applications S
PATI
AL
STR
UC
TUR
EYesHandles spatial structureNo
Spatial aggregate systems models e.g. ECOSPACE
Yes
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Ew
E
NO
. OF
SP
EC
IES
/ C
OM
PO
NE
NTS
30 +
20
10
532
ATLA
NTI
S
ER
SE
M,
INV
ITR
O,
OS
MO
SE
GA
DG
ET
BO
RM
ICO
N,
MS
VPA
, MU
LTS
PE
C,
MS
M,
SE
AP
OD
YM
ES
AM
, M
RM
, S
EA
STA
R,
Bio
ener
-ge
tic ty
pe
EwE, ATLANTIS, INVITRO
External forcing
Phytoplankton, detritus
Zooplankton, filter-feeders
Clupeoids, demersals etc
Marine mammals, sharks etc
TRO
PH
IC L
EV
EL
ERSEM SSEM
MRM, MSVPA, GADGET, SEASTAR, SEAPODYM, IBM, MSM, Bioenergetic
OSM
OSE
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Strategical Model Considerations and the Best Practice Approach
(based on report from the July 2007 FAO Workshop)
Consideration in Model Development
Best practice approach : ‘ideal’ practices, i.e. considerations when developing models. Not anticipated that these practices are always achievable or required.
Setting up a model
How many species or groups?
Aggregate based on shared characteristics of the species and omit the least important to keep web tractable
Include age, size or stage structure of the species of interest?
Include if there are major shifts over the course of the life history
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Modelling predator-prey interactions:
How much detail in representing predator-prey interactions?
Represent as bi-directional unless it can be motivated sufficiently strongly that it is adequate to include a one-way interaction only in which the predator ration is fixed and changes in prey abundance have no effect on predator populations
Which functional response?
Test sensitivity and robustness to alternative functional relationships
PREY DENSITY
PR
EY
KIL
LED
PE
R P
RE
DAT
OR
PE
R U
NIT
TIM
E
a) Type I
b) Type II
c) Type III
Predator abundance Bj
Tota
l con
sum
ptio
n ra
te Q
ij
(for f
ixed
pre
y ab
unda
nce
B i)
Bj (Input)
Default assumption is that thisis the present (input) situation
limit
limit/2
EwE Foraging arenaSUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Spatial considerations Best Practice ApproachInclude spatial structure?
Include where there are major shifts in the location of the species of interest over the course of its life history
Include seasonal and temporal structure?
Where there are large differences in the seasonal dynamics in species movement or production
Defining boundary conditions
Basing boundaries on biological rather than anthropogenic considerations such as national boundaries
Is fishery harvesting more than one stock of a particular species?
Model needs to distinguish such different stocks when the harvesting practice is such as might impact these stocks to different extents; this will necessitate spatially structured models
Distinguish different fleets?
If for the same mass of catch, they make substantially different impacts on target and bycatch species or on the habitat
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Model components Best Practice ApproachExplicitly represent primary productivity and nutrient cycling
May only be necessary when bottom-up forces or lower trophic levels are of key concern. Can be highly informative for some strategic modelling exercises.
Include environmental forcing?
Only if it is an absolute requirement for capturing system dynamics. When it is included there must be some means of generating future forcing for use in predictions and closed loop simulations.
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Model components
Best Practice Approach
How to model recruitment?
Recruitment may be included either as an emergent property or as a derived relationship Likely important for tactical and risk analyses, but not strict requirement for strategic models.
Mainly emergent recruitment
Mainly derived property
e.g. EwE, Atlantis, OSMOSE, GADGET
e.g. MSVPA, SEAPODYM, MSMSUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Model components Best Practice ApproachHow to model movement?
Testing sensitivity to a range of movement hypotheses. Parameterising movement matrices by fitting to these data. Decision rules check if resultant changes in distribution are sensible
EXAMPLES:OSMOSE: Spatially explicit with fish schools moving to areas with highest potential prey biomassGADGET: migration matrices specifying movement between areas; can parameterise by fitting to dataSEAPODYM: Movement model linked to habitat quality
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
External forcing Best Practice ApproachOther process error considerations
Other process error, arising from natural variation in model parameters, needs to be included when variation contributes substantially to uncertainty
Other anthropogenic forcing?
Their influence on shallow coastal and estuarine systems should be considered in conceptual models ; should be empirically included
Alternative stable states?
Strategic models in particular need to ensure forecasting the consequences of environmental change…
Explicitly consider fleet dynamics?
Important to consider if substantial changes to the spatial distribution of fishing may result from, for example, the declaration of an MPA.
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Technical and non-trophic:
Best Practice Approach
Technical interactions(e.g. multi-stock fisheries; by-catch)
If the bycaught species are themselves also subject to management, including stock rebuilding, or if the model aims to inform the level of bycatch of a threatened species.
Non-trophic interactions(e.g. habitat dependency and habitat mediated interactions and processes)
If a critical determinant of the dynamic of interest (e.g. biomass or abundance of a target group), or if management could be based around this interaction
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Dealing with uncertainty
Best Practice Approach
Should the model be fit to data?
Fitting to data is best practice, and this requires careful specification of likelihoods.
Taking account of parameter uncertainty
Include clear statements about uncertainties in model parameters; Bayesian methods and bootstrapping in ESAM and MRMs; …..
Model structure uncertainty
Identify alternative qualitative hypotheses for all of the processes considered likely to have an important impact on model outputs
What features to include in closed loop simulations?
Evaluation of feedback control harvest strategies should involve simulating the scheme that is likely to be actually used to determine management actions
Implementation uncertainty
Implementation uncertainty needs to be linked to consideration of fleet dynamics and is largely driven by, and must be included in, economic considerations…
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Use and outputs Best Practice ApproachShould code be freely available
Documentation and source code must be freely available to allow for review and understanding of the model. Using existing models can be of great help in learning, but careful thought is required when using a pre-existing model so that the tool is not misused
Social and economic outputs
Have economic experts collaborating with fisheries ecologists when designing a model implementation of economic factors
Ease of modularization
Best is object-oriented design
e.g. EwE, GADGET, SEPODYM
e.g. GADGET, new EwE
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Recent trends in model development
• Modularisation – e.g. substitute different growth, functional response modules
• Fitting to time series data• Computing constraints – e.g. running on multiple
computers in parallel using PVM • Spatial considerations• Representation of socio-economic factors and human
behavioural drivers• Multiple sector dynamics and management • Representation of biodiversity • Multi-species/ecosystem MSEs
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
OUTLINE OF TALKEcosystem Model Objectives Ecosystem Model Types Questions for EAF* Modelling Ecosystem Model Classification Considerations in Model Building and the
Best Practice Approach Role of Management Procedures Data requirements Conclusions
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Operating Model to simulate “true dynamics of resource
OPE
RAT
ING
M
OD
EL
Simulation testing
Explore uncertainties re model specification and fit to data
Methods and rules to compute Catch per ssmu
Use “future” data to compute Catch per ssmu
MA
NA
GEM
ENT
PRO
CED
UR
ERole of OMP/MP/MSE’s
(MP = Management procedure; MSE = Management Strategy Evaluation)
From Rademeyer et al. 2007SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Role of OMP/MP/MSE’s(MP = Management procedure; MSE = Management Strategy Evaluation)
• Approach involves an evaluation of the implications of alternative combinations of monitoring data, analytical procedures, and decision rules to provide advice on management measures that are robust to inherent uncertainties in all inputs and assumptions used.
• MSE or MP frameworks are used to identify and model uncertainties and to balance different resource dynamics representations.
• They provide key examples of formal methods for addressing uncertainty issues.
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
More re Dealing with Uncertainty• Few ecosystem models with applications to practical fisheries
management• Management Procedure testing procedures can use changes
in single species parameters (such as carrying capacity K) as a surrogate for ecological ecosystem effects e.g. climate change that are difficult to incorporate explicitly in operating models
• Technical ecosystem effects such as bycatch concerns can also be included as Robustness tests in the MP testing process
• These additions constitute a first step towards incorporating ecosystem aspects into practical fisheries management advice
• Multi-species/Ecosystem MPs being developed
Pg. 52SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Multi-species/Ecosystem MPs• ATLANTIS used to evaluate the performance of
ecological indicators • ATLANTIS used to test ecosystem models such as EwE
by generating simulated data with known parameters • South African Pelagic OMP - food requirements of
predators such as penguins need to be accounted for in the management process
• CCAMLR: FOOSA and SMOM – spatially explicit multi-species MP frameworks
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Spatial Multi-species Operating Model (SMOM) of Krill-Predator Interactions
Area 3 Feedback comparison
Peng
uin
num
bers
Seal
num
bers
0
50000
100000
150000
200000
250000
300000
2005
2010
2015
2020
2025
2030
2035
2040
2045
0.00
0.10
0.20
0.30
0.40
0.50
0.60
0.70
0.80
0.90
2005
2008
2011
2014
2017 2020
2023
0.00
0.10
0.20
0.30
0.40
0.50
0.60
0.70
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0.00
0.20
0.40
0.60
0.80
1.00
1.20
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0.00
0.50
1.00
1.50
2.00
2.50
3.00
3.50
4.00
2005
2008
2011
2014
2017 2020
2023
0.00
1.00
2.00
3.00
4.00
5.00
6.00
7.00
8.00
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0.00
0.20
0.40
0.60
0.80
1.00
1.20
1.40
1.60
1.80
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0
500
1000
1500
2000
2500
2005
2008
2011
2014
2017 2020
2023
0
100
200
300
400
500
600
700
800
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0
200
400
600
800
1000
1200
1400
1600
1800
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0
50
100
150
200
250
300
350
400
450
2005
2009
2013
2017
2021
0
100
200
300
400
500
600
700
800
900
1000
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0
50
100
150
200
250
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0
100
200
300
400
500
600
700
800
2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024
0100002000030000400005000060000
700008000090000
2005
2009
2013
2017
2021
2025
2029
2033
2037
2041
2045
0
200000
400000
600000
800000
1000000
1200000
1400000
0
2000
4000
6000
8000
10000
12000
14000
16000
2005
2009
2013
2017
2021
2025
2029
2033
2037
2041
2045
0
50
100
150
200
250
300
350
0
5000000000000
10000000000000
15000000000000
20000000000000
25000000000000
30000000000000
35000000000000
40000000000000
2005
2011
2017
2023
0
2000000000000
4000000000000
6000000000000
8000000000000
10000000000000
12000000000000
14000000000000
16000000000000
2005
2008
2011
2014
2017
2020
2023
2026
2029
2032
2035
2038
2041
2044
0
5000000000000
10000000000000
15000000000000
20000000000000
25000000000000
2005
2008
2011
2014
2017
2020
2023
2026
2029
2032
2035
2038
2041
2044
Area 10 Feedback comparison
0
200000
400000
600000
800000
1000000
1200000
1400000
1600000
1800000
2005
2008
2011
2014
2017
2020
2023
2026
2029
2032
2035
2038
2041
2044
0
500
1000
1500
2000
2500
3000
2005
2009
2013
2017
2021
2025
2029
2033
2037
2041
2045
0
100000
200000
300000
400000
500000
600000
0
200
400
600
800
1000
1200
1400
1600
2005
2008
2011
2014
2017
2020
2023
2026
2029
2032
2035
2038
2041
2044
0
5000000000000
10000000000000
15000000000000
20000000000000
25000000000000
30000000000000
35000000000000
40000000000000
2005
2007
2009
2011
2013
2015
2017
2019
2021
2023
2025
2027
2029
2031
2033
2035
2037
2039
2041
2043
2045
0
500000000000
1000000000000
1500000000000
2000000000000
2500000000000
2005
2007
2009
2011
2013
2015
2017
2019
2021
2023
2025
2027
2029
2031
2033
2035
2037
2039
2041
2043
2045
0
500000000000
1000000000000
1500000000000
2000000000000
2500000000000
3000000000000
3500000000000
4000000000000
2005
2007
2009
2011
2013
2015
2017
2019
2021
2023
2025
2027
2029
2031
2033
2035
2037
2039
2041
2043
2045
SMOM-predicted change in predator abundance with a) no feedback in spatial catch allocations and b) using a feedback control rule based on a moderate amount monitoring information available for all SSMUs.
No feedback
Feedback
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
Data Requirements
MRM DETAILED DATA RE FEW SPECIES – USUALLY SIZE/AGE STRUCTURE DATA
WHOLE ECO-
SYSTEM
LESS DATA RE MORE SPECIES
MORE DATA RE MORE SPECIES e.g. ADDING AGE STRUCTURE
SPATIAL
DATA RE MOVEMENT / DISTRIBUTIONS
ENVIRONMENT, ECONOMIC, SOCIAL, FISHING FLEET, ANTHROPOGENIC
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service
KESIMPULAN1. A good range of models have been developed for the task of
EAF, but greater focus is needed on strengthening these approaches and conducting the necessary data collection and experimentation to underpin confidence in these approaches
2. Management decisions will be enhanced by exploring the same issue with different models; confidence in the decisions will increase when the models independently converge on the same management decisions and when uncertainties in the results have been adequately considered.
3. MSE/MP approach is best practice4. Strategical modelling will mainly be used to inform and
evaluate the Ecosystem Approach to Fisheries, with use in tactical decisions rare
SUMBER: www.st.nmfs.noaa.gov/.../03_Review_...National Marine Fisheries Service