Responding to climate-driven changes in growth in the modern stock assessment models

Giancarlo M. Correa\(^1\), Cole Monnahan\(^2\), Jane Sullivan\(^3\), James T. Thorson\(^2\), Andre E. Punt\(^1\)

\(^1\)University of Washington, Seattle, WA

\(^2\)Alaska Fisheries Science Center, NOAA, Seattle, WA

\(^3\)Alaska Fisheries Science Center, NOAA, Juneau, AK

Somatic growth

Somatic growth


Growth in fishes

  • Stage 1: Energy to somatic growth
  • Stage 2: Energy to somatic growth and sexual maturation
  • Stage 3: Energy to reproduction

Morais and Bellwood (2020)

Somatic growth

Important contributor to biomass variability

Stawitz and Essington (2019)

Somatic growth

External factors that affect growth variability

Wilson et al. (2019)

State-space assessment models

State-space assessment models

Allows separation and estimation of both observation and process errors.


Observation equation: \(E[Y_t \mid X_t]= g(X_t,\theta)\)

Process equation: \(E[X_t \mid X_{t-1}] = h(X_{t-1},\theta)\)


\(X_t\) is the unobserved state at time step \(t\) and \(Y_t\) are observations. \(\theta\) is the vector of all unknown model parameters (fixed effects). \(X_t\) treated as random effects (Aeberhard et al. 2018).

State-space assessment models


Why use them? (Aeberhard et al. 2018, Stock and Miller, 2021)

  • handle structural breaks, time-varying parameters

  • handle missing observations

  • can be used to do forecasting

  • model complex and nonlinear relationships

  • less retrospective bias

The Woods Hole Assessment Model (WHAM)

  • Fully state-space age-structured model
  • Data: catch, indices of abundance, age compositions, empirical weight-at-age, environmental covariates
  • Separability of total catch and proportions-at-age, as well as annual F and selectivity
  • Random effects in selectivity, M, NAA, Q
  • Written in TMB and R (user friendly!) ( see R package ).

Stock and Miller (2021)

The Woods Hole Assessment Model (WHAM)


Limitations:

  • Lack of best practices for random effects use
  • growth cannot be modeled internally
  • selectivity-at-age options but not selectivity-at-length
  • single sex, no spatial component, no aging error

Growth modeling in state-space models

Data inputs



  • (Marginal) length compositions
  • Conditional age-at-length

Both are important sources of information to model growth internally.

Growth modeling


Growth modeling overview

In the population:

Mean length-at-age (LAA)

Growth equation (von Bertalanffy)


The mean length-at-age at the start of the year (\(y=1\)):

\[\tilde{L}_{y,a} = L_{\infty}+(L_1 - L_{\infty})exp(-k(a-1))\]

\(a=1\) is first age in WHAM. Then, when \(y>1\):

\[\tilde{L}_{y,a} = \begin{cases} L_1, & \mbox{if } a=1 \\ \tilde{L}_{y-1,a-1}+(\tilde{L}_{y-1,a-1}-L_{\infty})(exp(-k)-1) & \mbox{otherwise} \end{cases}\]

Mean length-at-age (LAA)

Growth equation (von Bertalanffy)


Then, to calculate the mean length-at-age at any fraction of the year:

\[L_{y,a} = \tilde{L}_{y,a} + (\tilde{L}_{y,a} - L_{\infty})(exp(-kf_y)-1)\] \(f_y\) is the year fraction.

Mean length-at-age (LAA)

Growth equation (von Bertalanffy)

Random effects on growth parameters can be predicted (notice log scale):

\[log(L_{\infty_t}) = \mu_{L_\infty} + \delta_{1,t}\]

\[log(k_t) = \mu_{k} + \delta_{2,t}\]

\[log(L_{1_t}) = \mu_{L_1} + \delta_{3,t}\]

\(t\) represents year or cohort effects and can be \(iid\) or \(AR1\).

Mean length-at-age (LAA)

LAA random effects

For this case, mean length-at-age ( \(\mu_{\tilde{L}_{a}}\), notice log scale ) are assumed to be parameters and can be estimated. \(\sigma_{y,a}\) still needed.

Time variability can be modeled by predicting random effects:

\[log(\tilde{L}_{y,a}) = \mu_{\tilde{L}_{a}} + \delta_{y,a}\]

\(\delta_{y,a}\) can be \(iid\) or \(2dAR1\) (full variance-covariance matrix).

Age-length transition matrix


Also, \(L_{y,a}\) and variation of length-at-age ( \(\sigma_{y,a}\) ) are used to calculate the age-length transition matrix (Stock Synthesis - SS - approach):

\[\varphi_{y,l,a} = \begin{cases} \Phi(\frac{L'_{min}-L_{y,a}}{\sigma_{y,a}}) & \mbox{for } l=1 \\ \Phi(\frac{L'_{l+1} - L_{y,a}}{\sigma_{y,a}}) - \Phi(\frac{L'_{l} - L_{y,a}}{\sigma_{y,a}}) & \mbox{for } 1<l<n_L \\ 1-\Phi(\frac{L'_{max} - L_{y,a}}{\sigma_{y,a}}) & \mbox{for } l = n_L \end{cases}\]

Where \(\Phi\) is standard normal cumulative density function, \(L'_{l}\) is the lower limit of length bin \(l\), \(L'_{min}\) is the upper limit of the smallest length bin, \(L'_{max}\) is the lower limit of the largest length bin, and \(n_L\) is the largest length bin index.

Age-length transition matrix

Mean weight-at-age (WAA)

Length-weight relationship

Optional when empirical weight-at-age not provided:

\[w_l = \Omega_1 l^{\Omega_2}\]

Random effects on \(\Omega_1\) and \(\Omega_2\) can also be predicted.

Then:

\[\hat{w}_{y,a} = \sum_l \varphi_{y,l,a}w_l\]

\(\hat{w}_{y,a}\) can also be fitted to \(w_{y,a}\) (observed mean weight-at-age)

Mean weight-at-age (WAA)

WAA random effects


Like the LAA random effects. Mean weight-at-age ( \(\mu_{\tilde{W}_{a}}\), notice log scale ) are assumed to be parameters and can be estimated.

Time variability can be modeled by predicting random effects:

\[log(\tilde{W}_{y,a}) = \mu_{\tilde{W}_{a}} + \delta_{y,a}\]

\(\delta_{y,a}\) can be \(iid\) or \(2dAR1\) (full variance-covariance matrix).

Selectivity

Originally, only selectivity-at-age functions were available (age-specific, logistic, double-logistic, decreasing-logistic)

New functions added:

  • double-normal: by age. SS-like (Methot and Wetzel, 2013)

  • len-logistic: increasing logistic at length

  • len-decreasing-logistic: by length

  • len-double-normal: by length

Environmental covariates


WHAM separates process (random walk or AR) and observation error for environmental covariates.

An environmental covariate can be linked to a state (i.e. parameter):

\[P_t = P exp(\beta_1 X_t)\]

\(P\) is the base state (parameter) value. Other links are also available (polynomials). Lags can be modeled.

Examples

Examples


Using SS ( ss3sim, Anderson et al., 2014 ), we simulated data that was then incorporated into WHAM.

  1. Growth variability
  2. Conditional age-at-length (CAAL) data
  3. Length-weight variability (condition factor)

Example 1: Growth variability


Simulated data:

  • catch (one fishery)
  • index of abundance (one survey)
  • marginal length compositions (fishery and survey)
  • temporal variability in asymptotic length ( \(L_\infty\) ) using PDO as the driver

Example 1: Growth variability


We implemented three models in WHAM:


  1. Random effects (iid_y) on \(L_\infty\)
  2. Random effects (iid) on mean length-at-age (LAA)
  3. Include an environmental variable (PDO), linked to \(L_\infty\)

Example 1: Growth variability

Example 2: CAAL data


Simulated data:

  • catch (one fishery)
  • index of abundance (one survey)
  • length compositions (fishery)
  • conditional age-at-length (survey)
  • selectivity-at-length (double-normal)

Example 2: CAAL data

CAAL residuals:

Example 2: CAAL data

Estimated selectivity-at-length:

Example 3: Length-weight variability


Simulated data:

  • catch (one fishery)
  • index of abundance (one survey)
  • length compositions (fishery and survey)
  • empirical weight-at-age data fitted
  • temporal variability in the \(\Omega_1\) parameter (L-W relationship)

Example 3: Length-weight variability

Walleye pollock in the Gulf of Alaska (GOA)

Model overview

Data:

  • One fishery (1970 to 2021) and six surveys
  • Age compositions for fishery and surveys
  • Mean weight-at-age for fishery and surveys

Parameters (penalized ML for time-varying quantities):

  • Time-varying fishery age selectivity
  • Two Qs vary over time
  • Recruitment variability

Starting model

ADMB model vs WHAM model

Growth variability

Observed survey fish lengths:

Growth modeling strategies


Model comparison

Mean SSB estimates:

Model comparison

SSB coefficient of variation:

Model comparison

Growth parameters (only for growth parametric approach):

Model comparison

Predicted mean length-at-age (Jan 1st) vs survey observations (\(\sim\) March 1st, not included in the model):

Model comparison

AIC values for models with same input data:


Model name (Marginal) AIC \(\Delta\) AIC
LAA random effects (\(iid\)) 816.7 0
vB equation (\(iid_y\)) 4036.6 3219.9
vB equation (\(iid_c\)) 4761.9 3945.2
WAA random effects (\(iid\)) 1177.2 360.5

Final thoughts


  • New tool to explore growth modeling in state-space assessment models
  • Non parametric approach seems to outperform a parametric equation
  • Non parametric approach is more flexible and deals with fish shrinkage

Future directions (for this project)


  • Include survey length information (e.g. marginal length compositions, conditional age-at-length data)
  • Add environmental covariate
  • Simulation experiment: compare strategies to account for growth variability using WHAM
    • Good practices for modeling growth in state-space models

Future directions (in general)


  • Software development: include sex, intraannual variability, tagging data, spatial structure
  • Comparison among platforms: e.g. SS vs WHAM?
  • Ecological research: meaning of all random effects options

Thanks

Cole Monnahan, Jane Sullivan, Jim Thorson, Andre Punt, Tim Miller, Jim Ianelli, Brian Stock

Find more information:
tinyurl.com/wham-growth