9.1 Fisheries Science
Fisheries science applies population ecology, stock assessment, and oceanographic knowledge to manage the sustainable harvest of marine fish and invertebrates. With global wild-capture fisheries yielding approximately 90 million tonnes per year and over one-third of assessed stocks classified as overfished, quantitative tools for fisheries management are essential for food security and marine conservation.
Logistic Population Model
The foundation of fisheries dynamics is the logistic growth equation, which describes how a population grows toward an environmental carrying capacity $K$. The intrinsic growth rate $r$ governs how quickly the population increases at low abundance, while the density-dependent term $(1 - N/K)$ slows growth as the population approaches $K$.
$$\frac{dN}{dt} = rN\left(1 - \frac{N}{K}\right)$$
$N$ = population size, $r$ = intrinsic growth rate, $K$ = carrying capacity
The surplus production is maximized when the population is at half its carrying capacity, i.e., $N = K/2$. At this point the maximum sustainable yield (MSY) is:
$$\text{MSY} = \frac{rK}{4}$$
Maximum sustainable yield occurs at $B_{\text{MSY}} = K/2$
~90 Mt/yr
Global wild capture
~34%
Stocks overfished
~60%
Maximally sustainably fished
Schaefer Surplus Production Model
The Schaefer model links biomass dynamics to fishing effort $E$. Catch is proportional to effort and biomass through the catchability coefficient $q$, so catch $C = qEB$. The biomass dynamics become:
$$\frac{dB}{dt} = rB\left(1 - \frac{B}{K}\right) - qEB$$
$q$ = catchability coefficient, $E$ = fishing effort
At equilibrium ($dB/dt = 0$), the equilibrium biomass is $B_{eq} = K(1 - qE/r)$ and the equilibrium yield is:
$$Y_{eq} = qEK\left(1 - \frac{qE}{r}\right)$$
Yield is a parabolic function of effort; the peak is MSY
The catch per unit effort (CPUE) serves as an index of abundance: $\text{CPUE} = C/E = qB$. Declining CPUE signals stock depletion and is a key warning indicator in fisheries management.
Stock-Recruitment Relationships
Stock-recruitment models relate the spawning stock biomass (SSB) to the number of recruits (young fish entering the fishery). The Beverton-Holt model assumes density-dependent juvenile mortality, while the Ricker model includes overcompensation at high stock sizes.
Beverton-Holt
$$R = \frac{\alpha S}{1 + \beta S}$$
Recruitment saturates at high stock; no overcompensation
Ricker
$$R = \alpha S \, e^{-\beta S}$$
Recruitment declines at very high stock due to cannibalism/competition
Here $S$ is the spawning stock biomass, $R$ is recruitment, and $\alpha$, $\beta$ are species-specific parameters estimated from data. The steepness parameter $h$ (recruitment at 20% of unfished SSB relative to unfished recruitment) is widely used to characterize productivity.
Fishing Mortality & Overfishing
The total instantaneous mortality rate $Z$ is the sum of natural mortality $M$ and fishing mortality $F$:
$$Z = F + M, \quad N(t) = N_0 \, e^{-Zt}$$
Exponential decline in numbers-at-age under constant mortality
Growth Overfishing
Fish caught before reaching optimal harvest size; yield per recruit is reduced
Recruitment Overfishing
SSB reduced below level needed for adequate recruitment; stock collapse risk
Ecosystem Overfishing
Removal of top predators alters food web structure; "fishing down the food web"
The exploitation rate $\mu = F/Z \cdot (1 - e^{-Z})$ gives the fraction of the population removed by fishing each year. The reference point $F_{\text{MSY}}$ is the fishing mortality that produces maximum sustainable yield. Overfishing occurs when $F > F_{\text{MSY}}$.
Fisheries Management Tools
Total Allowable Catch (TAC)
Science-based annual catch limits set from stock assessments. Divided among participants via national quotas or individual allocations.
Individual Transferable Quotas (ITQ)
Market-based allocation of fishing rights. Reduces "race to fish" and overcapitalization. Tradeable shares of TAC.
Marine Protected Areas (MPAs)
No-take zones allow stock recovery. Spillover benefits adjacent fishing grounds. Larval export supports recruitment.
Ecosystem-Based Management (EBM)
Considers entire food webs, habitat, bycatch, and environmental variability rather than single-species targets. Precautionary approach.
Bycatch—the incidental capture of non-target species—remains a major challenge. Turtle excluder devices (TEDs), circle hooks, and acoustic deterrents help reduce bycatch of marine mammals, seabirds, and sea turtles.
Python: Logistic Growth with Harvesting & Stock-Recruitment
Python: Logistic Growth with Harvesting & Stock-Recruitment
Python!/usr/bin/env python3
Click Run to execute the Python code
Code will be executed with Python 3 on the server
Fortran: Age-Structured Virtual Population Analysis (VPA)
Virtual Population Analysis reconstructs historical population numbers-at-age from catch-at-age data. Working backwards from terminal estimates, VPA solves the catch equation$C_{a,t} = \frac{F_{a,t}}{Z_{a,t}} N_{a,t} (1 - e^{-Z_{a,t}})$ for $F_{a,t}$ and $N_{a,t}$.
Fortran: Age-Structured Virtual Population Analysis (VPA)
FortranAge-structured VPA (cohort analysis) for fish stock assessment
Click Run to execute the Fortran code
Code will be compiled with gfortran and executed on the server
Yield Per Recruit Analysis
The Beverton-Holt yield-per-recruit (YPR) model determines the optimal fishing mortality and age at first capture. The yield per recruit is:
$$\frac{Y}{R} = F \, e^{-M(t_c - t_r)} \sum_{n=0}^{3} \frac{U_n \, e^{-n K_g (t_c - t_0)}}{F + M + n K_g} \left(1 - e^{-(F+M+nK_g)(t_\lambda - t_c)}\right)$$
$t_c$ = age at first capture, $t_r$ = age at recruitment,$K_g$ = von Bertalanffy growth parameter
$F_{0.1}$ Reference Point
The fishing mortality where the slope of the YPR curve is 10% of its value at the origin. A more conservative target than $F_{\text{max}}$.
$F_{\text{max}}$ Reference Point
The fishing mortality that maximizes yield per recruit. Often leads to recruitment overfishing and is not recommended as a management target.
Derivation: Logistic Growth & MSY (Schaefer Model)
Step 1: Logistic Growth Equation
Begin with a population of size $B$ growing at intrinsic rate $r$ toward carrying capacity $K$. The density-dependent growth is:
$$\frac{dB}{dt} = rB\left(1 - \frac{B}{K}\right)$$
Step 2: Introduce Fishing as a Harvest Term
In the Schaefer model, the catch rate is proportional to both fishing effort $E$ and biomass $B$ through catchability $q$: $C = qEB$. The biomass dynamics with harvest become:
$$\frac{dB}{dt} = rB\left(1 - \frac{B}{K}\right) - qEB$$
Step 3: Find Equilibrium Biomass
At equilibrium, $dB/dt = 0$. Factor out $B$ (discarding $B = 0$):
$$r\left(1 - \frac{B_{eq}}{K}\right) = qE \quad \Longrightarrow \quad B_{eq} = K\left(1 - \frac{qE}{r}\right)$$
Step 4: Derive Equilibrium Yield
Substitute $B_{eq}$ into $Y_{eq} = qEB_{eq}$:
$$Y_{eq} = qEK\left(1 - \frac{qE}{r}\right)$$
Step 5: Maximize Yield to Find MSY
Differentiate $Y_{eq}$ with respect to $E$ and set the derivative to zero:
$$\frac{dY_{eq}}{dE} = qK - \frac{2q^2KE}{r} = 0 \quad \Longrightarrow \quad E_{\text{MSY}} = \frac{r}{2q}$$
Step 6: Compute MSY and $B_{\text{MSY}}$
Substitute $E_{\text{MSY}}$ back into the equilibrium biomass and yield expressions:
$$B_{\text{MSY}} = K\left(1 - \frac{q \cdot r/(2q)}{r}\right) = \frac{K}{2}$$
$$\text{MSY} = qE_{\text{MSY}} \cdot B_{\text{MSY}} = q \cdot \frac{r}{2q} \cdot \frac{K}{2} = \frac{rK}{4}$$
Step 7: CPUE as Abundance Index
The catch per unit effort (CPUE) is directly proportional to biomass:
$$\text{CPUE} = \frac{C}{E} = \frac{qEB}{E} = qB$$
Thus CPUE serves as a relative index of stock abundance; linear regression of CPUE on effort estimates $q$ and $K$.
Step 8: Fishing Mortality at MSY
Define fishing mortality as $F = qE$. At MSY:
$$F_{\text{MSY}} = qE_{\text{MSY}} = q \cdot \frac{r}{2q} = \frac{r}{2}$$
Overfishing occurs when $F > F_{\text{MSY}} = r/2$. The stock is overfished when $B < B_{\text{MSY}} = K/2$.
Derivation: Beverton-Holt Stock-Recruitment Model
Step 1: Density-Dependent Juvenile Mortality Assumption
Assume that juvenile survival decreases linearly with population density. The instantaneous mortality rate for juveniles is $\mu(N) = \mu_0 + \mu_1 N$, where $N$ is the number of juveniles competing for resources.
$$\frac{dN}{dt} = -(\mu_0 + \mu_1 N) N$$
Step 2: Solve the Differential Equation
This is a Bernoulli equation. Let $u = 1/N$, so $du/dt = -N^{-2}\,dN/dt$. Substituting:
$$\frac{du}{dt} = \mu_0 u + \mu_1$$
This linear ODE has the solution:
$$u(t) = \left(u_0 + \frac{\mu_1}{\mu_0}\right)e^{\mu_0 t} - \frac{\mu_1}{\mu_0}$$
Step 3: Derive the Recruitment Function
After the juvenile period of duration $\tau$, recruitment $R = N(\tau) = 1/u(\tau)$. With the initial number of eggs proportional to spawning stock $S$: $N(0) = \gamma S$, we obtain:
$$R = \frac{\gamma S \, e^{-\mu_0 \tau}}{1 + \frac{\mu_1 \gamma S}{\mu_0}(1 - e^{-\mu_0 \tau})}$$
Step 4: Standard Beverton-Holt Form
Define $\alpha = \gamma e^{-\mu_0 \tau}$ and $\beta = \frac{\mu_1 \gamma}{\mu_0}(1 - e^{-\mu_0 \tau})$ to obtain the classic form:
$$R = \frac{\alpha S}{1 + \beta S}$$
Step 5: Properties of the Beverton-Holt Curve
At low stock sizes ($S \to 0$): $R \approx \alpha S$ (linear increase). The slope at the origin is the maximum recruits-per-spawner $\alpha$.
$$\lim_{S \to \infty} R = \frac{\alpha}{\beta} \quad \text{(asymptotic maximum recruitment)}$$
Step 6: Steepness Parameterization
Define steepness $h$ as the fraction of unfished recruitment $R_0$ produced when SSB is 20% of unfished SSB $S_0$:
$$h = \frac{R(0.2 S_0)}{R_0} = \frac{\alpha \cdot 0.2 S_0}{(1 + \beta \cdot 0.2 S_0) R_0}$$
Steepness ranges from 0.2 (strong density dependence) to 1.0 (recruitment independent of stock). High steepness means the stock is resilient to fishing.
Derivation: Surplus Production Model Fitting
Step 1: Discrete Biomass Dynamics
Discretize the Schaefer model over annual time steps. The biomass next year equals this year's biomass plus surplus production minus catch:
$$B_{t+1} = B_t + rB_t\left(1 - \frac{B_t}{K}\right) - C_t$$
Step 2: Link CPUE to Biomass
The observed CPUE index $I_t$ is assumed proportional to biomass: $I_t = qB_t$. This allows expressing the equilibrium yield in terms of observables:
$$Y_{eq} = \frac{r}{q} I\left(1 - \frac{I}{qK}\right)$$
Step 3: Observation Model and Likelihood
Assume log-normal observation errors in the CPUE index. The likelihood of the data given parameters $\theta = (r, K, q, \sigma)$ is:
$$\mathcal{L}(\theta | \text{data}) = \prod_{t=1}^{T} \frac{1}{I_t \sigma \sqrt{2\pi}} \exp\left(-\frac{(\ln I_t - \ln qB_t)^2}{2\sigma^2}\right)$$
Step 4: Negative Log-Likelihood
Taking the negative log-likelihood and dropping constants:
$$-\ln \mathcal{L} = \frac{T}{2}\ln\sigma^2 + \frac{1}{2\sigma^2}\sum_{t=1}^{T}(\ln I_t - \ln q - \ln B_t)^2$$
Step 5: Concentrate Out $q$ and $\sigma^2$
The MLE for $q$ is the geometric mean ratio of CPUE to predicted biomass. Define residuals $\epsilon_t = \ln I_t - \ln B_t$:
$$\hat{q} = \exp\left(\frac{1}{T}\sum_{t=1}^{T} \epsilon_t\right), \quad \hat{\sigma}^2 = \frac{1}{T}\sum_{t=1}^{T}(\epsilon_t - \bar{\epsilon})^2$$
Step 6: Management Quantities from Fitted Model
Once $r$ and $K$ are estimated, all management reference points follow directly:
$$\text{MSY} = \frac{rK}{4}, \quad B_{\text{MSY}} = \frac{K}{2}, \quad F_{\text{MSY}} = \frac{r}{2}, \quad E_{\text{MSY}} = \frac{r}{2q}$$
The current stock status is assessed by comparing $B_{\text{current}}/B_{\text{MSY}}$ and $F_{\text{current}}/F_{\text{MSY}}$ on a Kobe plot.
Key Equations Summary
Baranov Catch Equation
$$C = \frac{F}{Z} N_0 \left(1 - e^{-Z}\right)$$
Exploitation Rate
$$\mu = \frac{F}{Z}\left(1 - e^{-Z}\right) = \frac{C}{N_0}$$
Spawning Potential Ratio (SPR)
$$\text{SPR} = \frac{\text{SSB/R under fishing}}{\text{SSB/R unfished}}$$
Target: SPR > 0.30–0.40 to prevent recruitment overfishing