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Alt text: Fisheries models range from single and multispecies models to full ecosystem models

Georges Bank multispecies model
keyrun project

Overview
ICES WGSAM review, 10 and 14 October 2022

Sarah Gaichas, Andy Beet, Kiersten Curti, Gavin Fay, Robert Gamble,
Ron Klasky, Sean Lucey, Maria Cristina Perez, and Howard Townsend

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Alt text: Fisheries models range from single and multispecies models to full ecosystem models

Why include species interactions?

Ignore predation at your peril: results from multispecies state-space modeling (Trijoulet, et al., 2020a)

Ignoring trophic interactions that occur in marine ecosystems induces bias in stock assessment outputs and results in low model predictive ability with subsequently biased reference points.

VanessaPaper

EM1: multispecies state space

EM2: multispecies, no process error

EM3: single sp. state space, constant M

EM4: single sp. state space, age-varying M

note difference in scale of bias for single species!

modcomp

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This is an important paper both because it demonstrates the importance of addressing strong species interactions, and it shows that measures of fit do not indicate good model predictive performance. Ignoring process error caused bias, but much smaller than ignoring species interactions. See also Vanessa's earlier paper evaluating diet data interactions with multispecies models

EBFM Objectives in the Northeast US

  • EBFM Objective 1: what happens with all the species in the region under a certain management regime?

    • Apply a full system model to assess "side effects" of target species management
    • Ability to implement fishing and biological scenarios
    • Hypothesis testing and MSE framework desirable
  • EBFM Objective 2: how well do multispecies models perform for assessment?

    • Consider alternative model structures
      • Biomass dynamics
      • Size structured
      • Age structured
    • Evaluate data availability for each structure
    • Evaluate estimation performance of each structure
    • Evaluate uncertainty and sensitivity
    • Evaluate feasibility of developing and using multi-model inference
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MS-Keyrun model development and testing objectives are based on general ecosystem based management questions as well as specific discussions regarding EBFM development in New England. We will use this as an opportunity to address questions about the effects of management on the broader ecosystem, and about performance of assessment tools.

Place-based approach

"Place-based" means a common spatial footprint based on ecological production, which contrasts with the current species-based management system of stock-defined spatial footprints that differ by stock and species.

The medium blue area in the map is Georges Bank as defined by NEFSC trawl survey strata. SOE = State of the Ecosystem report

The input data for this project differs from the input data for most current stock assessments, and the results of these multispecies assessments are not directly comparable with current single species assessments.

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The project currently implements several place-based multispecies assessment models and one food web model. "Place-based" means a common spatial footprint based on ecological production, which contrasts with the current species-based management system of stock-defined spatial footprints that differ by stock and species. (See stock area comparisons.) Therefore, the input data for this project differs from the input data for most current stock assessments, and the results of these multispecies assessments are not directly comparable with current single species assessments. However, similar processes can be applied to evaluate these models. Georges Bank as defined for this project uses the NEFSC bottom trawl survey strata highlighted in medium blue below, which corresponds to the spatial unit for survey-derived ecosystem indicators in the Northeast Fisheries Science Center (NEFSC) New England State of the Ecosystem (SOE) report. Orange outlines indicate the ten minute square definitions for Ecological Production Units defined by a previous analysis.

Objective 1: evaluate system responses to management

Rpath (Lucey, et al., 2020a) with MSE capability (Lucey, et al., 2021)

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Food web: Rpath in collaboration with AFSC

Species interactions:

  • Full predator-prey: Consumption leads to prey mortality and predator growth
  • Static and dynamic model components

Static model: For each group, ii, specify:

Biomass BB (or Ecotrophic Efficiency EEEE)
Population growth rate PBPB
Consumption rate QBQB
Diet composition DCDC
Fishery catch CC
Biomass accumulation BABA
Im/emigration IMIM and EMEM

Solving for EEEE (or BB) for each group:

Bi(PB)iEEi+IMi+BAi=j[Bj(QB)jDCij]+EMi+CiBi(PB)iEEi+IMi+BAi=j[Bj(QB)jDCij]+EMi+Ci

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Predation mortality M2ij=DCijQBjBjBiM2ij=DCijQBjBjBi

Fishing mortality Fi=ng=1(Cig,land+Cig,disc)BiFi=ng=1(Cig,land+Cig,disc)Bi

Other mortality M0i=PBi(1EEi)

Food web: Rpath in collaboration with AFSC

Dynamic model (with MSE capability):

dBidt=(1AiUi)jQ(Bi,Bj)jQ(Bj,Bi)M0iBiCmBi Consumption:

Q(Bi,Bj)=Qij(VijYpredjVij1+(1Sij)Ypredj+Sik(αkjYpredk))×(DijYpreyθijiDij1+((1Hij)Ypreyi+Hik(βikYpreyk))θij)
Where Vij is vulnerability, Dij is “handling time” accounting for predator saturation, and Y is relative biomass which may be modified by a foraging time multiplier Ftime,

Y[pred|prey]j=FtimejBjBj

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The parameters Sij and Hij are flags that control whether the predator density dependence Sij or prey density dependence Hij are affected solely by the biomass levels of the particular predator and prey, or whether a suite of other species’ biomasses in similar roles impact the relationship.

For the default value for Sij of 0 (off), the predator density dependence is only a function of that predator biomass and likewise for prey with the default value of 0 for Hij.

Values greater than 0 allow for a density-dependent effects to be affected by a weighted sum across all species for predators, and for prey. The weights αkj and βkj are normalized such that the sum for each functional response (i.e. kαkj and kβkj for the functional response between predator j and prey i) sum to 1. The weights are calculated from the density-independent search rates for each predator/prey pair, which is equal to 2QijVij/(Vij1)BiBj.

Objective 2: evaluate multispecies assessment tools

Multispecies production (Gamble, et al., 2009a)

Multispecies catch at length (Gaichas, et al., 2017a)

(and eventually)

Multispecies catch at age (Curti, et al., 2013a)

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Multispecies production simulation: Kraken and estimation FIT: MSSPM

Species interactions:

  • Predation: Top down (predation decreases population growth of prey, predator population growth independent of prey)
  • Competition: Within and between species groups

Based on Shaefer and Lotka-Volterra population dynamics and predation equations

  • Species have intrinsic population growth rate ri
  • Full model has
    • Carrying capacity K at the species group level KG and at the full system level Kσ
    • Within group competition βig and between group competition βiG slow population growth near K
    • Predation αip and harvest Hi reduce population

dNidt=riNi(1NiKGgβigNgKGGβiGNGKσKG)NipαipNpHiNi

  • Simpler version used in most applications has interaction coefficient α that incorporates carrying capacity Bi,t+1=Bi,t+riBi,tBi,tjαi,jBj,tCi,t
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Interaction coefficients αi,j can be positive or negative

C can be a Catch time series, an exploitation rate time series Bi,tFi,t or an qE (catchability/Effort) time series.

Environmental covariates can be included on growth or carrying capacity (in the model forms that have an explicit carrying capacity).

Multispecies catch at length simulation model: Hydra

Species interactions:

  • Predation: Top down only (predators increase M of prey, predators grow regardless of prey)

Based on standard structured stock assessment population dynamics equations, Same MSVPA predation equation as MSCAA (but length based), same dependencies and caveats

  • First, split M for species i size j into components: Mi,j,t=M1i+M2i,j,t
  • Calculate M2 with MSVPA predation equation, which applies a predator consumption:biomass ratio to the suitable prey biomass for that predator.
  • Suitability, ρ, of prey species m size n for a given predator species i size j a function of size preference and vulnerability {0,1}.
  • Food intake I for each predator-at-size is temperature dependent consumption rate times mean stomach content weight.
  • Also sensitive to "other food" Ω.

M2m,n,t=ijIi,j,tNi,j,tρi,j,m,nabρi,j,a,bWa,bNa,b+Ω

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But:

  • Covariates on growth, maturity, recruitment possible; intended for environmental variables
  • So could hack in prey-dependent growth but making it dynamic is difficult

We specify 'preferred' predator-prey weight ratio (log scale) Ψj and variance in predator size preference σj to compare with the actual predator-prey weight ratio (wn/wj) to get the size preference ϑ.

ϑn,j=1(wn/wj)σj2πe[loge(wn/wj)Ψj]2σ2j

Food intake is Ii,j,t=24[δjeωiT]ˉCi,j,k,t

Model life cycle

(NRC, 2007)

What makes a good model?

  • Differs by life stage

  • Each builds on the next

  • Common themes

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This week: constructed model (framework) review

For each model, reviews should evaluate:

  1. Spatial and temporal resolution
  2. Algorithm choices
  3. Assumptions (scientific basis, computational infrastructure; adequacy of conceptual model)
  4. Data availability/software tools
  5. Quality assurance/quality control (code testing)
  6. Test scenarios
  7. Corroboration with observations
  8. Uncertainty/sensitivity analysis
  9. Peer review (previous)
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Common attributes across models

A common dataset for 10 Georges Bank species has been developed, as well as a simulated dataset for model performance testing. The mskeyrun data package holds both datasets. All modeling teams used these datasets. Group decisions on data are also documented online.

Years: 1968-2019

Area: Georges Bank (previous map)

Species:
Atlantic cod (Gadus morhua),
Atlantic herring (Clupea harengus),
Atlantic mackerel (Scomber scombrus),
Goosefish (Lophius americanus),
Haddock (Melanogrammus aeglefinus),
Silver hake (Merluccius bilinearis),
Spiny dogfish (Squalus acanthias),
Winter flounder (Pseudopleuronectes americanus),
Winter skate (Leucoraja ocellata), and
Yellowtail flounder (Limanda ferruginea)

foodweb

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Datasets: real and simulated

Real data from NEFSC databases via R packages survdat, comlandr, mscatch

mscatchlogo

Simulated data from Norwegian Barents Sea Atlantis model via R package atlantisom

Norwegian-Barents Sea

Hansen et al. 2016, 2018

NOBA

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ms-keyrun real diet data

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Modelers discuss overview of results

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References

Curti, K. L. et al. (2013a). "Evaluating the performance of a multispecies statistical catch-at-age model". En. In: Canadian Journal of Fisheries and Aquatic Sciences 70.3, pp. 470-484. ISSN: 0706-652X, 1205-7533. DOI: 10.1139/cjfas-2012-0229. URL: http://www.nrcresearchpress.com/doi/abs/10.1139/cjfas-2012-0229 (visited on Jan. 13, 2016).

Gaichas, S. K. et al. (2017a). "Combining stock, multispecies, and ecosystem level fishery objectives within an operational management procedure: simulations to start the conversation". In: ICES Journal of Marine Science 74.2, pp. 552-565. ISSN: 1054-3139. DOI: 10.1093/icesjms/fsw119. URL: https://academic.oup.com/icesjms/article/74/2/552/2669545/Combining-stock-multispecies-and-ecosystem-level (visited on Oct. 18, 2017).

Gamble, R. J. et al. (2009a). "Analyzing the tradeoffs among ecological and fishing effects on an example fish community: A multispecies (fisheries) production model". En. In: Ecological Modelling 220.19, pp. 2570-2582. ISSN: 03043800. DOI: 10.1016/j.ecolmodel.2009.06.022. URL: http://linkinghub.elsevier.com/retrieve/pii/S0304380009003998 (visited on Oct. 13, 2016).

Lucey, S. M. et al. (2021). "Evaluating fishery management strategies using an ecosystem model as an operating model". En. In: Fisheries Research 234, p. 105780. ISSN: 0165-7836. DOI: 10.1016/j.fishres.2020.105780. URL: http://www.sciencedirect.com/science/article/pii/S0165783620302976 (visited on Dec. 09, 2020).

Lucey, S. M. et al. (2020a). "Conducting reproducible ecosystem modeling using the open source mass balance model Rpath". En. In: Ecological Modelling 427, p. 109057. ISSN: 0304-3800. DOI: 10.1016/j.ecolmodel.2020.109057. URL: http://www.sciencedirect.com/science/article/pii/S0304380020301290 (visited on Apr. 27, 2020).

NRC (2007). "Chapter 4. Model Evaluation". En. In: Models in Environmental Regulatory Decision Making. Washington D.C.: The National Academies Press, pp. 104-169. DOI: 10.17226/11972. URL: https://www.nap.edu/read/11972/chapter/6 (visited on Aug. 29, 2019).

Trijoulet, V. et al. (2020a). "Performance of a state-space multispecies model: What are the consequences of ignoring predation and process errors in stock assessments?" En. In: Journal of Applied Ecology n/a.n/a. ISSN: 1365-2664. DOI: 10.1111/1365-2664.13515. URL: https://besjournals.onlinelibrary.wiley.com/doi/abs/10.1111/1365-2664.13515 (visited on Dec. 04, 2019).

Additional resources

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Hydra: past use as simulation model

2018 CIE for Ecosystem Based Fishery Management Strategy

hydrascenarios

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Next: Multispecies catch at age estimation model: seeking catchy name FIT: MSCAA

Species interactions:

  • Predation: Top down only (predators increase M of prey, predators grow regardless of prey)

Based on standard age structured stock assessment population dynamics equations

  • First, split M for species i age a into components: Mi,a,t=M1i+M2i,a,t
  • Calculate M2 with MSVPA predation equation, which applies a predator consumption:biomass ratio to the suitable prey biomass for that predator.
  • Suitability is a function of predator size preference (based on an age-specific predator:prey weight ratio) and prey vulnerability (everything about the prey that isn't size related).
  • Also sensitive to "other food"

M2i,a,t=1Ni,a,tWi,a,tjbCBj,bBj,b,tϕi,a,j,b,tϕj,b,t

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Size preference is gi,a,j,b,t=exp[12σ2i,j(lnWj,b,tWi,a,tηi,j)2]

Suitability, ν of prey i to predator j: νi,a,j,b,t=ρi,jgi,a,j,b,t

Scaled suitability: ˜νi,a,j,b,t=νi,a,j,b,tiaνi,a,j,b,t+νother

Suitable biomass of prey i to predator j: ϕi,a,j,b,t=˜νi,a,j,b,tBi,a,t

Available biomass of other food, where Bother is system biomass minus modeled species biomass: ϕother=˜νotherBother,t Total available prey biomass: ϕj,b,t=ϕother+iaϕi,a,j,b,t

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Alt text: Fisheries models range from single and multispecies models to full ecosystem models

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