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03_marlin_paper.qmd
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---
format:
pdf:
toc: false
fig-pos: 'H'
mainfont: Times New Roman
sansfont: Times New Roman
fontsize: 12pt
keep-tex: true
fig-format: png
html: default
docx:
reference-doc: template.docx
execute:
echo: false
message: false
warning: false
cache: false
csl: apa.csl
bibliography: references.bib
header-includes:
- \usepackage{setspace}\doublespacing
- \usepackage{lineno}\linenumbers
- \usepackage{bm}
editor:
markdown:
wrap: 80
---
```{r setup}
#| echo: false
#| message: false
#| include: false
source(here::here("00_setup.R"))
theme_set(theme_classic(base_size = 10, base_family = "Helvetica") + theme(strip.background = element_blank()))
if (!file.exists(here("data","sci_to_com.csv"))){
sci_to_com <- taxize::sci2comm(unique(marlin_inputs$scientific_name),simplify= TRUE,db='ncbi') %>%
bind_rows(.id = "critter") %>%
pivot_longer(everything(),names_to = "critter",values_to = "common_name") %>%
mutate(common_name = stringr::str_to_title(common_name))
write_csv(sci_to_com, file = here("data","sci_to_com.csv"))
} else{
sci_to_com <- read_csv(file = here("data","sci_to_com.csv"))
}
print(results_path)
results_files <- list.files(file.path(results_path))
rdses <- results_files[str_detect(results_files,'.rds')]
walk(rdses,~assign(str_remove(.x,".rds"), read_rds(file.path(results_path,.x)), envir = .GlobalEnv)) #load and assign all .rds objects in the appropriate results folder
small_width <- 80 # figure width in mm
big_width <- 180 # width in mm
```
## Title Page
Title: Simulating Benefits, Costs, and Trade-offs of Spatial Management in
Marine Social-Ecological Systems
Running Title: Spatial Social-Ecological Simulation
Author Names: Daniel Ovando^1,2^, Darcy Bradley^3,4,5^, Echelle
Burns^3,4,5^,Lennon Thomas^3,4,5^, James Thorson^6^
Affiliations: ^1^ School of Aquatic and Fishery Sciences, University of
Washington,1122 NE Boat St Box 355020,Seattle, WA, USA ^3^ Marine Science
Institute, University of California, Santa Barbara, CA, USA ^4^ Bren School of
Environmental Science & Management, University of California, Santa Barbara, CA,
USA ^5^ Environmental Markets Lab, University of California, Santa Barbara, CA,
USA ^6^ Habitat and Ecological Processes Research Program, Alaska Fisheries
Science Center, NOAA, NMFS, NOAA, Seattle, WA, USA
Corresponding Author: Daniel Ovando
[dovando\@iattc.org](mailto:[email protected]){.email}
Present Address: \^2 Inter-American Tropical Tuna Commission,8901 La Jolla
Shores Drive, La Jolla, CA 92037, USA
Author Contributions: Methods and code written by DO and JT. Results produced by
DO. Data collected by EB and LT. All authors contributed to the
conceptualization and writing of the manuscript.
Conflict of Interest Statement: The authors declare no conflicts of interest.
\newpage
## Abstract
Designing effective spatial management strategies is challenging because marine
ecosystems are highly dynamic and opaque, and extractive entities such as
fishing fleets respond endogenously to ecosystem changes in ways that depend
upon ecological and policy context. We present a modeling framework, `marlin`,
that can be used to efficiently simulate the bio-economic dynamics of marine
systems in support of both management and research. We demonstrate `marlin’s`
capabilities by focusing on two case studies on the conservation and food
production impacts of marine protected areas (MPAs): a coastal coral reef and a
pelagic tuna fishery. In the coastal coral reef example, we explore how
heterogeneity in species distributions and fleet preferences can affect
distributional outcomes of MPAs. In the pelagic case study, we show how our
model can be used to assess the climate resilience of different MPA design
strategies, as well as the climate sensitivity of different fishing fleets. This
paper demonstrates how intermediate complexity simulation of coupled
bio-economic dynamics can help communities predict and potentially manage
trade-offs between conservation, fisheries yields, and distributional outcomes
of management policies affected by spatial bio-economic dynamics.
## Keywords
Bio-economic modeling; Conservation planning; Fisheries; Marine protected areas; Marine spatial planning; Spatio-temporal modeling
## Table of Contents
1. Introduction
2. Methods
1. Model Summary
2. Fleet Dynamics
3. Population Model
4. Case Studies
3. Results
1. Coral Reefs
2. Pelagic Systems
4. Discussion
1. Insights from Case Studies
2. Putting `marlin` in Context
3. General Recommendations for Use
5. Acknowledgements
6. Data Availability Statement
7. References
8. Figure Legends
\newpage
## Introduction
Communities around the world are increasingly looking to spatial strategies for
managing marine ecosystems. For example, the "30x30" movement calls for
implementation of marine protected areas (MPAs), a form of spatial management,
across 30% of the world's oceans by the year 2030 [@grorud-colvert2021 and
references therein]. Recent agreements such as the United Nations (UN) agreement
on the conservation and sustainable use of marine biodiversity of areas beyond
national jurisdiction (BBNJ) calls for expanded area-based management in the
high seas. However, designing spatial management strategies to achieve desired
objectives -- which may include recovery and resilience of overfished species,
increased food production and economic well being, and equitable distribution of
benefits - is not straightforward. Different species have different resilience
to fishing, distributions in space and time, and value to fishing fleets;
fishing fleets themselves can have varying ranges of incentives and reactions to
policy structures. These dynamics pose challenges even for single-species
assessment and management strategies, which are only amplified when we consider
management policies designed around multiple species, fleets, and
spatial features in the oceans [@field2006a].
To illustrate, policies such as MPAs designed for both conservation and food
production must consider factors such as the optimal size and placement of a
protected area network given a wide array of life history, species
distributions, exploitation levels, fleet dynamics, and policy-dependent
behavior [@reimer2017], all of which may fluctuate over time, particularly given
the impacts of climate change. Efforts to effectively design spatial management
strategies such as MPAs are further constrained by a lack of empirical evidence
describing the size [@ovando2021a] and time required [@nickols2019] for MPAs to
produce substantial effects across a range of social-ecological settings
[@mcclanahan2021].
Many models have been developed to support the theory and design of spatial
management strategies (see @fulton2015 for a review).
However, these models have tended to be either highly complex and tactical
models designed for use in a specific location, or to be extremely stylized
representations intended to provide theoretical insights with less relevance for
specific applications. This lack of accessible models capable of representing reasonable amounts of complexity presents a challenge to stakeholders charged with marine spatial
planning exercises. It also presents a barrier to the scientific community,
wherein comparisons across spatial management simulation studies are clouded by
discrepancies in underlying model structures beyond differences in the core
phenomenon in question.
To help address this challenge, we present a bio-economic modeling tool
called [`marlin`](https://danovando.github.io/marlin/) that allows for efficient
simulation of spatio-temporal biological and economic dynamics. `marlin` allows
users to simulate the impacts of marine management policies across a range of
species targeted by various fleets across heterogeneous and dynamic seascapes.
This model can help users seek Pareto-optimal solutions that produce win-wins or
minimize trade-offs across multiple management objectives, such as food
production and conservation [@lester2013].
Here we present the core methods and functionality of `marlin`, and demonstrate
its use in two case-study applications; a coral reef system and a pelagic
system. Our results show how interacting ecological, economic, and design
attributes can affect the degree of benefits, costs, and trade-offs between
conservation and food production outcomes of MPAs. More broadly, this paper
demonstrates the functionality of our model and the critical need for
considering sufficiently realistic coupled ecological and economic dynamics in
policy evaluations.
## Methods
The model presented here focuses on representing the dynamics of heterogeneous
habitats and movement dynamics, along with the simultaneous impacts of fishing
fleets across multiple species (frequently called "technical interactions"). In
its general form, `marlin` simulates the behavior of populations of spatially
explicit biologically independent animal populations with age and subsequent
size structure affected by one or more fishing fleets in time steps and a
spatial resolution specified by the user.
The front end of the software package accompanying this paper,
[`marlin`](https://danovando.github.io/marlin/), is written in R
[@rcoreteam2021] to facilitate use, while the underlying population model is
written in C++, integrated through the `Rcpp` package [@eddelbuettel2018]. Users on standard computers should be able to simulate one age-structured population distributed across two-dimensional spatial surface represented by a
ten by ten grid of cells over 20 years in fractions of a second. It is very important to note that the parameters of
this model cannot be fit to data directly within the `marlin` package; users
must set model parameters themselves based on externally available data and
their best judgement.
While `marlin` does not simulate species interactions, fishing fleets in
`marlin` are capable of targeting and affecting multiple species simultaneously.
This allows the model to simulate processes such as fisheries bycatch or effort
displacement in a way that accounts for how fleet behavior might affect multiple
species in the system. The ability to track the impacts of fishing fleets across
multiple species simultaneously is particularly important as very few fishing
fleets are truly single species.
We define a few commonly used terms here. *Yields* refers to the volume of catch
from fishing activity. *Spawning stock biomass* refers to the total biomass of
reproductively mature fish in the water, a function of numbers, weight,
fecundity, and sexual maturity at age. We measure the size of the population by
the ratio of the *spawning stock biomass* (SSB) in a given time step relative to
*unfished spawning stock biomass* (SSB0), SSB/SSB0. An SSB/SSB0 value of 1 means
that the population is unfished, a SSB/SSB0 value of 0 means the population is
extinct.
As a demonstration of the use of this modeling tool, we explore two general
situations
1. Trade-offs and distributional outcomes for food production and multi-species
biomass in a coastal coral reef fishery
2. Implications of climate-driven range shifts for MPA design in a pelagic
fishery
We chose these two examples to illustrate the use of `marlin` in contrasting
systems in which spatial management strategies such as MPAs are increasingly
considered.
Below we provide a summary of the `marlin` model, as well as details of the case
studies.
### Model Summary
`marlin` simulates the dynamics of one or more species, currently best
representing fish-like species, using age-structured population dynamics. Ages
are then converted to lengths using the von Bertalanffy growth equation with
log-normally distributed variation in the length at age. Each time step, species
move throughout the simulated seascape using both diffusion and "taxis" (active
movement towards preferred habitat), experience natural and potentially fishing
mortality, and potentially spawn using one of the possible forms of
density dependence implemented in the model, with the ability to include
auto-correlated stochastic deviations in the amount of offspring produced,
generally called "recruitment deviates".
These species can be caught by fishing fleets. A fishing fleet is defined in
`marlin` by a set of fishers that have the same fishing skill, prices, and
contact selectivity [@sampson2014] for individual species (each of which we
denote as a *métier*). For example, both a longline and purse-seine fleet may
capture bigeye (*Thunnus obesus*, Scombridae) and skipjack (*Katsuwonus pelamis*, Scombridae) tunas, but the longline fleet can
be made much more likely to catch larger bigeye than skipjack, and *vice versa*.
It is important to note that contact selectivity reflects the ability of the
fishing method in question to capture fish of different sizes that come into
"contact" with the gear. The contact selectivity of each of the specified
*métiers* will then interact with the distribution of fish sizes and fishing
effort in space simulated by `marlin` to produce a net "population" selectivity,
which may differ from the individual contact selectivities of each of the
*métiers* [@waterhouse2014; @sampson2014]. Each fishing fleet distributes its
fishing effort in space according to a specified spatial allocation rule (see
*Spatial Allocation of Effort* section), for example in proportion to
profit-per-unit-effort, conditional on management policies in places such as
quotas and/or the presence of any spatial restrictions such as MPAs.
Unconstrained by management, the total amount of effort exerted by each fleet
can follow one of two dynamics: open access or constant effort (see *Calculating Total Effort*). Under constant effort, the total amount of effort of each fleet
is fixed over time, with the possible exception of attrition due to MPA
placement. Under open access, the total amount of effort in a given time step is
a function of the profitability of the fleet in the previous time step, evolving
until a bionomic equilibrium of zero total profits is reached. Profitability is
a function of the volume and price of each species caught, as well as the cost
of the total amount of fishing effort per fleet and the travel costs as a
function of distance from a port (see *Fleet Dynamics*).
The dynamics of each fleet can be modified by management in a variety of ways.
Managers can impose size limits for individual species within each fleet. They
can also impose total catch quotas for each species in each time step. When
quotas are activated, if the total catch across all fleets for a given species
under the model's effort dynamics would exceed the allowable quota for that
species, the total effort for each fleet is decreased proportional to its
contribution to the total catch until the quota is satisfied. As an alternative
to catch quotas, managers can set an effort cap per fleet, which prevents effort
from exceeding a given amount under open access dynamics, though effort may be
reduced below this cap if required by the profit equation (i.e. the fleet can
choose to fish less than the quota or effort cap). `marlin` also allows users to specify closed fishing seasons for one or more
species in the system. Lastly, managers can specify locations of no-take MPAs, which
can change in size and location if needed. When MPAs are implemented, fishing
effort that used to operate inside the MPAs can either leave the fishery, or be
redistributed outside the MPA (the default behavior).
On the biological side, at a minimum users must for each species being simulated supply the common or scientific name of the species in question, a
measure of the level of fishing intensity the species is experiencing at the
start of the simulation, and the diffusion rates for adult and larvae. Given a
common or scientific name, the model will then input default life history
parameters based on FishLife [internet connection required, @thorson2020],
though users are encouraged to check these values and supply their own life
history parameter values when possible.
On the fleet side, users must for each fleet they wish to simulate at a minimum
specify the contact selectivity curves for each species caught by the fleet, as
well relative price per unit weights greater than 0 for any target species.
Species caught by but not targeted by the fleet can be represented by prices of
0 (not targeted) or below 0 (actively avoided).
### Fleet Dynamics
Each fishing fleet (*f*) generates catches or yield (*Y*), revenues (*R*), and
costs (*C*) from fishing individual species (*s*) that it targets in a given
time step (*t*) and patch (*p*). The fleet then makes decisions around fishing
locations and intensity, subject to regulatory constraints, based on the total
profitability across all species in space and time.
Revenues for each fleet in a time step are a function of the total amount of
each species caught and its price ($\Pi$). The amount caught is a function of
the contact selectivity at age of each species for each fleet ($\alpha$), the
fishing efficiency of the fleet for that that species (also called
"catchability", *q*), the amount of fishing effort of the fleet in question in
that patch in a given time step (*E*), and the total instantaneous fishing
mortality (*u*) at age (*a*) (including all other fleets) for that particular
species in that patch and time step.
$$
R_{t,p,f} = \sum_s^{N_s} \sum_a^{N_a} \Pi_{s,f}\frac{\alpha_{a,s,f}q_{s,f}E_{t,p,f}}{u_{t,p,s,a}} \times Y_{t,p,s,a}
$$ {#eq-revenue}
Total catches or yield *Y* are calculated through the Baranov equation
[@baranov1918], which accounts for the total instantaneous mortality (both
fishing and natural, *z*) and divides the total amount of the population biomass
(*b*) killed between natural (*m*) and fishing (*u*) sources, with the amount of
biomass killed through fishing called "yield".
$$
Y_{t,p,s,a} = \frac{u_{t,p,s,a}}{z_{t,p,s,a}} \times b_{t,p,s,a} \times (1 - e^{-z_{t,p,s,a}})
$$ {#eq-yield}
$$
u_{t,p,s,a} = \sum_{f=1}^{N_f}\alpha_{a,s,f}q_{s,f}E_{t,p,f}
$$ {#eq-u}
Contact selectivity at age $\alpha$ is modeled through either a logistic form or
a dome-shaped form. The logistic form is based on the lengths *l* at which 50%
of individuals are selected by the fishing gear, $l^{sel}$, and $\delta$ which
is the difference between the length at 50% selectivity and the length at 95%
selectivity.
$$
\alpha_{a,s,f}=\frac{1}{(1 + e^{-log(19)\times\frac{l_{a,s} - l^{sel}_{s,f}}{\delta^{sel}_{s,f}}})}
$$ {#eq-selectivity}
We approximate the dome-shaped form as a normal distribution with mean $l^{sel}$
and standard deviation $\sigma$. The normal density function is re-scaled such
that the selectivity is 1 at $l^{sel}$.
$$
\alpha_{a,s,f}= \frac{1}{\sigma_{s,f} \sqrt{2\pi}}e^{-0.5 \left( \frac{l_{a,s} - l^{sel}}{\sigma_{s,f}} \right)^2}
$$ {#eq-dome}
Lastly *z* is total mortality, the sum of fishing mortality (*u*) and natural
mortality (*m*).
$$
z_{t,p,s,a} = u_{t,p,s,a} + m_{s,a}
$$ {#eq-z}
This means that each species experiences a total mortality at age in a time step
in a patch, individual fractions of which are portioned off as catches and
subsequently revenues for each fleet. Given revenues *R*, we then calculate
profits $\phi$ based on revenues and costs. Costs *C* are calculated as a
function of base costs per unit effort ($\gamma$) as well as potential
additional costs per unit effort of fishing in particular patches ($\eta$) for
fleet *f*. $\beta$ allows for the cost per unit effort effort to scale
non-linearly. Travel costs ($\eta$) are calculated based on the Euclidean
distance of each patch to the nearest port of a given fleet, and users can
specify any number from zero to the number of patches of port locations. When no
ports are specified, travel costs are zero.
$$
\phi_{t,p,f} = R_{t,p,f} - C_{t,p,f}
$$ {#eq-profits}
$$
C_{t,p,f} = \gamma_f \left( E_{t,p,f}^{\beta_f} + \eta_{f,p} E_{t,p,f} \right)
$$ {#eq-travel-costs}
#### Calculating Total Effort
`marlin` allows for two general modes of effort. The simplest is "constant
effort", in which the total effort of each fishing fleet remains constant over
time.
The more complex option is "open access". Under open access, the total amount of
effort in a given time step for fleet *f* is a function of the profitability of
that fleet in the proceeding time steps in which fishing was open, where
$\theta$ controls the responsiveness of fleet *f* to the log of the ratio of
total revenues *R* to total costs *C*, and approximates the proportional change
in effort in response to a one unit changes in the log revenue to cost ratio.
$$
E_{t+1,f} = E_{t,f} \times e^{\theta_f log(R_{t,f} / C_{t,f})}
$$ {#eq-oa}
This is essentially a Gompertz model for fishing effort, that has been used for
other theoretical studies of fishing effort dynamics [@thorson2013].
#### Tuning Fishing Fleet Parameters
The degree of fishing pressure exerted by a given set of fishing fleets on each
species is a function of a range of parameters including the total amount of
effort (*E*), the contact selectivity ogives ($\alpha$), fishing cost (*C*), the spatial distribution of the
species affected by the fleet, the relative prices across species ($\Pi$), and
the catchability coefficient of each fleet for each species ($q$). `marlin`
provides a tuning option to help users achieve desired biological outcomes from
their fleets.
Users can tune their fleets in one of two ways, conditional on the underlying
population dynamics of the species in question. First, they can specify a target
exploitation rate *u* for each species in their simulation. Taking all the other
parameters of the model as given, `marlin` then adjusts the catchability
coefficients $q_{f,s}$ for each fleet *f* and species *s* such that the
equilibrium exploitation rate for each species matches the desired level.
Second, they can specify a target total spawning stock biomass under fishing
divided by total unfished spawning stock biomass, and the model will adjust the
catchability coefficients $q_{f,s}$ for each fleet *f* and species *s* such that
the equilibrium ratio of fished to unfished spawning biomass for each species
matches the desired level.
Users can also tune the fleet dynamics by specifying the ratio of costs to
revenues. Price data for use in the model can be obtained relatively simply
through literature reviews, market surveys, or local experts. However, cost
parameters are more complicated, as translating say cost per day of fishing into
the same representation of fishing effort used in the model is not straight
forward to accomplish. As an alternative, users can specify an equilibrium
cost-to-revenue ratio for the fleet, essentially the profit margins of the fleet
in question, and `marlin` will tune the cost parameter to achieve this desired
cost to revenue ratio given the other parameters in the model.
#### Spatial Allocation of Effort
Each fishing fleet decides how to allocate its effort in space based on one of
four possible spatial allocation strategies. The ideal free distribution (IFD)
is the standard method for distributing fishing fleets in spaces (see
@gillis2003 and references therein). However, analytical solutions to the IFD
present a number of complications for our model. The IFD for a single fleet
would commonly be modeled as a Nash-equilibrium based on the actions of each of
the individual fleet conditional on the actions of all other fleets. While
possible to implement, this would slow down our model runs to the point of
practically preventing large-scale evaluation of the spatial management
policies.
As such we explored a series of "next best" fleet distribution algorithms. While
not the IFD in any individual time step, over time they start to approximate the
IFD, as each assumes that fleets base their decisions for the current time step
on the outcomes in the prior timestep, meaning that the impacts of the actions
of other fishing fleets are eventually accounted for. The timeline required for the
fleets to reach an equilibrium distribution will vary and depend on factors such
as the population dynamics of the species in question and the number and degree of
competition across fleets. Users should explore different simulation times to
ensure that any results they wish to use are not simply a reflection of the
fleet dynamics fluctuating on their way to an equilibrium condition.
The four possible fleet distribution algorithms are
1. Revenue per unit effort (RPUE): The fleet distributes itself in space based
on the realized revenue per unit effort in each patch in the preceding time
step
2. Revenue: The fleet distributes itself in space based on the realized total
revenue in each patch in the preceding time step
3. Profit per unit effort (PPUE): The fleet distributes itself in space based
on the realized profit per unit effort in each patch in the preceding time
step
4. Profit: The fleet distributes itself in space based on profit in each patch
in the preceding time step
Revenue based spatial distribution is not likely to be very realistic; in
general we would expect fishing fleets to respond to profits on some core level.
However, due to the complexity of parameterizing cost functions, fleet dynamics
are often evaluated based on yield or revenue alone, and so we include those
scenarios here to allow users to evaluate the potential implications of this
choice. The decision on whether to allocate the fleet based on absolute or
relative (per unit effort) metrics is more complex. When effort represents the
actions of separate and individual fishing actors (e.g. independent fishing
vessels), a per-unit-effort strategy, in which fishers distribute themselves
based on the expected catch of their individual efforts, may be more realistic
[@hilborn1987]. Conversely, a system defined by a sole owner seeking to maximize
total profits might be better represented by a fleet model based on total
profits.
### Population Model
The underlying population model used is an age structured single-species model
in the manner of @ovando2021a. The population model requires many parameters.
However, if the user supplies either a scientific or common name for the species
in question (scientific preferred), the model will supply default values for
that species based on the values reported in FishLife [@thorson2020]. FishLife provides estimates of core life history
parameters for fish species based on a model integrating published values and
evolutionary connections. FishLife provides reasonable default values for given
species, but these default values should be not be taken as definitive and users
should check the default values and input best available parameters specific to the stock in question if they
wish to best represent the dynamics of their specific system.
#### Movement
`marlin`'s movement dynamics are based on a continuous-time Markov chain (CTMC),
as described in @thorson2021a. Within this framework, the model allows for
movement to be broken down into three components of *advection* (drifting with
currents), *taxis* (active movement towards preferred habitat), and *diffusion*
(essentially remaining variation in movement not explained by advection or
taxis). For now, `marlin` focuses just on the diffusion and taxis components of
this model, assuming that advection is zero, though future extensions could
incorporate advection vectors from oceanographic models. In this way, `marlin`
allows users to run anything from a simple Gaussian dispersal kernel up to a
system governed by species that passively diffuse out from a core habitat
defined by a dynamic thermal range. The model currently focuses on diffusion and
taxis, which allows for general representation of animals following physical or
oceanographic features, but research on the incorporation and importance of
advective forces would be of value going forward.
We provide a brief overview of the the general CTMC method here (see
@thorson2021a for a detailed description). Under this framework, movement of
individuals from each patch to each other patch in the system in a given
timestep *t* for life stage *a* of species *s* is defined by a movement matrix
$\pmb{M}_{t,s,a}$. $\pmb{M}_{t,s,a}$ is calculated as a function of diffusion
$\pmb{D}$ and taxis $\pmb{\tau}$ matrices scaled by the width of the time step
(e.g. one year) $\Delta_{t}$ and the length of the edge of each patch (e.g. one
kilometer) $\Delta_d$ specified by the user. This parameterization allows users
to set the effective area of the spatial domain through two avenues; the number
of patches, which effectively scales the resolution of the model, and the area
of each patch, which scales the spatial extent of the simulation.
The individual components (*M*) of the movement matrix ($\pmb{M}$) are filled
based on an adjacency matrix, which defines whether two patches are both
adjacent and water (as opposed to land or another physical barrier), a diffusion
rate $D$ defined in units of area of a patch per unit of time, and a habitat
preference function *H* in units of length of a side of a patch per unit time.
For example, if we are defining the time units as years and the distance units
as kilometers, for a tuna $D$ might be 1,000 $\frac{KM^2}{Year}$. We then use
parameters $\Delta_{t}$ and $\Delta_{d}$ parameters to translate the diffusion
rate $D$ to match the time step and patch size used in a simulation. For
example, if we were to run a model on a monthly timestep given time units of
years, then $\Delta_{t} = 1/12 years$. If one square patch in the simulation has
an area of 100km^2^, then $\Delta_d = 10KM$. This "scale free" parameterization
means that appropriate value of $D$ can be identified for a species and then
set, regardless of the time step or patch size used in the simulation model
itself.
The taxis component of the movement process is a function of the difference in
habitat quality *H*. The habitat preference function itself can take any form
the user wishes. Exponentiating the difference in the habitat preference
function between patches turns the taxis matrix into a multiplier of the
diffusion rate *D*. As such, when creating habitat layers for simulation, users
can tune the scale of the habitat gradient function to result in realistic
multipliers of the diffusion rate. This parameterization ensures that the
off-diagonal elements of the movement matrix $\pmb{M}_{t,s,a}$ are all
non-negative, a requirement of the CTMC method.
$$
M_{p1,p2,t,s,a} = \begin{cases}
= \frac{\Delta_{t}}{\Delta_{d}^2}De^{\frac{\Delta_t(H(p2,t,s,a) - H(p1,t,s,a))}{\Delta_d}} & \text{if p2 and p1 are adjacent}\\
= -\sum_{p' \neq p1} M_{p1,p2,t,s,a} & \text{if p1 = p2}\\
= 0 & \text{otherwise.}
\end{cases}
$$ {#eq-diffusion}
For both the diffusion and taxis matrices, we allow for the inclusion of
physical barriers to movement (i.e. land). Pairs of patches that are adjacent
but in which one or both patches are a barrier to movement are set as
non-adjacent. The CTMC model then produces movement dynamics that move around
barriers rather than over them.
The movement of individuals across patches is then calculated by matrix
multiplication of the pre-movement vector of the number of individuals
($\pmb{n}$) of species *s* at age *a* in time step *t* across all patches *p*
times the matrix exponential of the movement matrix $\pmb{M}$
$$
\pmb{n}_{t+1,s,a} = \pmb{n}_{t,s,a}e^{\pmb{M}_{t,s,a}}
$$ {#eq-movement}
While this CTMC approach to movement simulation is to date not commonly seen in
the marine modeling literature, it has numerous advantages that warrant its
seeming complexity. First, the parameters of the model have interpretable
biological meaning (e.g. the diffusion rate *D*). Second, when only diffusion is
present, the model will generalize to the familiar dynamics of a Gaussian
dispersal kernel at whatever spatial and temporal resolution the simulation is
set to. Third, the taxis model allow for clearly parameterized active habitat
choices by species, allowing us to simulate preferences of species in space and
time efficiently. Lastly, the CTMC form has the advantage that its parameters
are directly estimable from real data. So, if provided with for example spatial
abundance data and a tagging study, users can estimate the diffusion and taxis
movement parameters in the same manner as @thorson2021a, and then pass their
estimated parameters to `marlin` for simulation (so long as the estimating
method uses the same functional form as the movement model in `marlin`).
#### Population Growth
For the population model, numbers *N* at time *t* for age *a* are a function of
growth, death, and recruitment
$$
N_{t,p,s,a} = \begin{cases}
= BH(SSB_{t-1,p,s,a}) & \text{if $a = 1$}\\
= N_{t-1,p,s,a-1}e^{-(z_{t-1,p,s,a-1})}, & \text{if $1< a < max(age)$}\\
= N_{t-1,p,s,a}e^{-(z_{t-1,p,s,a})} + N_{t-1,a-1}e^{-(z_{t-1,p,s,a-1})}, & \text{if $a = max(a)$}
\end{cases}
$$ {#eq-pop}
where *BH* is the Beverton-Holt recruitment function [@beverton1957] and *SSB*
is spawning-stock-biomass. Per convention, the model allows for a "plus group",
wherein rather than tracking numbers of every possible age, individuals greater
than or equal to a given maximum age are grouped together.
Spawning stock biomass *SSB* is calculated by converting age to mean length at
age, calculating weight at age, maturity at age, and then calculating spawning
stock biomass as the sum of spawning potential at age in a given time step,
taking into account the potential for hyperallometry in the manner of
@marshall2021. Age is converted to length through the von Bertalanffy growth
equation given parameters asymptotic length ($l_\infty$), growth (*k*) and
theoretical age at length zero ($a0$) assuming log-normally distributed
variation *u* in the length at age with CV $\sigma_s$.
$$
l_{a,s} = l_{\infty,s}\left(1 - e^{-k_s(a - a0_{s})}\right)e^{u_s}
$$ {#eq-len}
$$
u_s \sim N(0,\sigma_s)
$$ {#eq-sigma}
Users can manually supply a vector of of natural mortality at age (*m*). Or,
they can supply one value of natural mortality which is then converted into
mortality at age through one of two means. Under the default behavior, natural
mortality at age given a target mean mortality across all ages $m_s$ is
calculated using a length-inverse mortality function [@lorenzen2022].
$$
minv_{s_a} = (\frac{l_{s,a}}{l_{\infty,s}})^{-1}
$$ {#eq-minv}
$$
m_{s,a} = \frac{minv_{s,a}}{mean( minv_{s,a})}m_s
$$ {#eq-mata}
Alternatively, users can set mortality at age to be constant
$$
m_{s,a} = m_s
$$ {#eq-mata2}
Biomass *B* at age is then given by the weight at length equation governed by a
scaling coefficient $\Omega_s$ and an exponent $\Phi_s$ that controls the rate
at which volume scales with length
$$
B_{a,s} = _s \times l_{a,s}^{wb_s}
$$ {#eq-weight}
The proportion of sexually mature individuals (*mat*) at a given age is then
calculated as a logistic function where $l_{mat}$ is the length at which on
average 50% of individuals are sexually mature, and $\delta_{mat}$ is the unit
of length beyond $l_{mat}$ at which on average 95% of fish are sexually mature.
$$
mat_{a,s} = \frac{1}{\left(1 + e^{-log(19)\times\frac{l_{a,s} - lmat_s}{{\delta}mat_s}}\right)}
$$ {#eq-mat)}
Spawning stock biomass at time *t* is then calculated as a function of the
numbers at age, the maturity at age, and the weight at age raised by a parameter
$\gamma$. When $\gamma$ is greater than 1, the species is said to experience
hyperallometric fecundity, i.e. fecundity increases faster than weight.
$$
SSB_{t,p,s} = \sum_{a=1}^{N_a}w_{a,s,t}^{\gamma_s} mat_{t,a} N_{t,p,s,a}
$$ {#eq-ssb}
#### Recruitment
Recruitment (i.e. the number of age 1 individuals entering the population)
follows Beverton-Holt dynamics parameterized around steepness (*h*) with
log-normally distributed recruitment deviates $\epsilon$. When steepness is one
recruitment is independent of spawning biomass. As steepness approaches 0.2
recruitment becomes a linear function of spawning biomass. `marlin` allows users
to specify a target unfinished spawning stock biomass ($SSB0$), which will be
achieved by tuning the total unfished recruitment ($r0$), given the remaining
life history parameters and independent of any characteristics of the fishing
fleets.
We allow for five variants in the timing of density dependent recruitment,
building off of @babcock2011 :
1. Global density dependence: Density dependent recruitment is a function of
the sum of spawning biomass across all patches, and recruits are then
distributed according to habitat quality
$$
\begin{aligned}
N_{t,p,s,a = 1} = \left(\frac{0.8{\times}\sum_{p=1}^P{r0_{p,s}}\times{h_s}\times\sum_{p=1}^P{SSB_{t-1,p,s}}}{0.2\times{\sum_{p=1}^PSSB0_{p,s}} \times(1 - h_s)+(h_s - 0.2) \times{\sum_{p=1}^PSSB_{t-1,p,s}}}\right) \times \\ {r0_{p,s} / \sum_p^P{r0_{p,s}}\times \epsilon_{t,s}
}
\end{aligned}
$$ {#eq-dd2}
where $r0$ is is a vector of recruits under unfished conditions in a given
patch.
2. Local density dependence: Density dependent recruitment occurs independently
in each patch and recruits are retained in their home patch.
$$
n_{t,p,s,a = 1} = \left(\frac{0.8{\times}r0_{p,s}\times{h_s}\times{SSB_{t-1,p,s}}}{0.2\times{SSB0_{p,s}}\times(1 - h_s)+(h_s - 0.2)\times{SSB_{t-1,p,s}}}\right) \times \epsilon_{t,s}
$$ {#eq-dd1}
3. Local density dependence then disperse: Density dependent recruitment occurs
independently in each patch and recruits are then dispersed.
$$
n_{t,p,s,a = 1} = \left(\frac{0.8{\times}r0_{p,s}\times{h_s}\times{SSB_{t-1,p,s}}}{0.2\times{SSB0_{p,s}}\times(1 - h_s)+(h_s - 0.2)\times{SSB_{t-1,p,s}}}\right)\times \pmb{d^l_s} \times \epsilon_{t,s}
$$ {#eq-dd3}
where **d^l^** is the recruitment movement matrix
4. Post-dispersal density dependence: Larvae are distributed throughout the
system, and then density dependent recruitment occurs based on the density
of spawning biomass at the destination patch.
$$
larv_{t,p,s} = SSB_{t-1,p,s} \times\pmb{d^l_s}
$$ {#eq-larvmove}
$$
n_{t,a = 1,p,s} = \left(\frac{0.8{\times}r0_{p,s}\times{h_s}\times{larv_{t,p,s}}}{0.2\times{SSB0_{p,s}}\times(1 - h_s)+(h_s - 0.2)\times{larv_{t,p,s}}}\right) \times \epsilon_{t,s}
$$ {#eq-dd4}
5. Global density dependence allocated by spawning biomass: Density dependence
is a function of the sum of spawning biomass across all patches, and
recruits are then distributed according to the distribution of spawning
biomass
$$
\begin{aligned}
n_{t,p,s,a = 1} = \left(\frac{0.8{\times}\sum_{p=1}^P{r0_{p,s}}\times{h_s}\times\sum_{p=1}^P{SSB_{t-1,p,s}}}{0.2\times{\sum_{p=1}^PSSB0_{p,s}}\times(1 - h_s)+(h_s - 0.2)\times{\sum_{p=1}^PSSB_{t-1,p,s}}}\right) \times \\ \frac{SSB_{t-1,p,s}}{\sum_{p=1}^PSSB_{t-1,p,s}} \times \epsilon_{t,s}
\end{aligned}
$$ {#eq-dd5}
Log-normal recruitment deviates are calculated with the potential for
autocorrelation defined with strength $\rho$
$$
\upsilon_{t,s} \sim \begin{cases}
N(0,\sigma_{r,s}), & \text{if $t = 1$}\\
\rho_s \upsilon_{t-1,s} + \sqrt{1 - \rho_s^2} N(0,\sigma_{r,s}), & \text{if $t > 1$}
\end{cases}
$$ {#eq-recdev}
And log recruitment deviates are converted to raw units using the bias
correction factor
$$
\epsilon_{t,s} = e^{\upsilon_{t,s} - \sigma_{r,s}^2/2}
$$ {#eq-bias}
#### Reference Points
Fisheries management is often concerned with measuring stock status relative to
maximum sustainable yield (MSY) based reference points, though the exact level
of stock status relative to MSY reference points desired by societal objectives may vary widely. MSY based reference points present a
problem for a multi-fleet and spatial-temporal model such as `marlin`. MSY and
the fishing mortality rate that would produce MSY, $F_{MSY}$, are a function of
fishery selectivity. Fishery selectivity in this model can vary by fleet, and
species can be distributed unevenly in both space and time. This means that the
net effective fishing selectivity on a species can vary depending on the
dynamics at a given moment, making the definition of an equilibrium concept such
as MSY challenging [@berger2017 and references therein].
As such, we do not report MSY based reference points in the model by default.
There are many different strategies for estimating reference points in spatially
explicit systems [@kapur2021a]. We leave it to users to define and find relevant
reference points as required by their specific needs.
### Case Studies
We include two examples demonstrating how `marlin` can be used to support marine
spatial planning. In the first, we show how `marlin` can be used to compare the
total and distributional impacts of MPAs designed in a heavily fished coastal
coral reef ecosystem. In the second, we demonstrate how `marlin` can be used to
assess components of climate resilience of alternative MPA design strategies in
a pelagic system.
Each of the case studies contains too many parameters and options to be
succinctly presented in the text here. Readers should consult the accompanying code to view
the precise details of each simulation. Targeted applications must carefully
consider and document all decisions made around model parameters.
#### MPA Design Strategies
We make use of three potential "rule of thumb" MPA design strategies in our case
studies
1. *Rate*: MPAs are placed based on the pre-MPA SSB/SSB0 weighted catch
relative to the total catch in a patch. So, patches with high rates of catch
of depleted species relative to total catch are prioritized.
2. *Target Fishing*: MPAs locations are prioritized proportional to fishery
catches. Patches with high total catches are prioritized over patches with
low catches.
3. *Spawning Ground*: MPAs are centered on the grounds of a known spawning
aggregation. This strategy is only used in the coral reef case study.
In theory, the design of MPA networks can be optimized through the use of a
modeling framework, and depending on the validity of the model, this process is
likely to produce better outcomes than manually-designed strategies
[@rassweiler2012; @rassweiler2014]. However, designing optimal MPA networks
becomes increasingly difficult as the range of objectives and the complexity of
the model increase. Therefore, we focus here on the design and performance of these more rule of thumb design
strategies that may be more accessible to a wider range of users. We allow all
MPAs to be designed in a mosaic fashion in these examples, but users can easily
extend the analysis to compare outcomes between contiguous (MPA is made up of
one continuously connected block) and mosaic (MPAs can be separated in space)
MPA designs [@pons2022].
#### Coastal Coral Reef Fishery
In our coastal coral reef example, we model the dynamics of four tropical
Pacific species: a grouper (*Epinephelus fuscoguttatus*, Serranidae), a shallow-reef snapper
(*Lutjanus malabaricus*, Lutjanidae), a deep-reef snapper (*Pristipomoides filamentosus*, Lutjanidae),
and a reef shark (*Carcharhinus amblyrhynchos*, Carcharhinidae). The simulated groupers undergo
a mass migration to a spawning aggregation once per year, followed by the
sharks. Shallow-reef snappers stay in reefs closer to shore above a steep
drop-off year-round, while deep-water snappers stay in the deeper reefs past the
drop-off (@fig-coral).
These species are targeted by two different fleets. Fleet One primarily targets
the grouper and near-shore snapper populations, but will land any incidentally
captured sharks. Fleet One has a logistic selectivity pattern for all species,
as they retain any fish caught for consumption or sale. Fleet One is totally
dependent on fishing for their livelihood, meaning the local community takes
advantage of every possible opportunity to fish, and as such we model it as a
"constant effort" fishery. Due to having less efficient boats, Fleet One has a
higher cost per distance coefficient than Fleet Two. Fleet One's home port is
located near the site of the grouper spawning grounds.
Fleet Two is a more commercial fleet that primarily targets the snapper
populations. This fleet primarily sells their catch to local restaurants and
distributors where plate-sized fish are prized, and so for both snapper and
grouper Fleet Two has a dome-shaped selectivity pattern [@kindsvater2017]. While
plate-sized deep snapper are the primary target of Fleet Two, we model Fleet
Two's selectivity for deep snapper as logistic due to high levels of discard
mortality for deep-water snapper resulting from barotrauma. Fleet Two catches
groupers, though less than Fleet One, and receives no price for sharks due to
the requirements of a certification program through which they sell their
deep-water snapper. Accidental captures (bycatch) of sharks do occur, which
results in mortality. Fleet Two operates under open-access dynamics, as fishing
is not the only means of subsistence for this community; short-term effort
expands and contracts in response to profitability of the primarily
grouper-driven fishery. Fleet two's home port is located in the northwest corner
of the simulation space.
We used `marlin` to simulate the outcomes for both food production and
conservation for each of the species and both of the fleets as a function of
both MPA size and MPA design strategy. For this exercise, MPAs are placed with
perfect information and have no design constraints for continuity. We ran the
simulation for in quarter year time steps for 20 years ($\Delta_t = 1/4$) and
set the area of each patch to be 5km^2^ ($\Delta_p = \sqrt{5}$), using 144
patches for a total area of 2,000 KM^2^.
#### Pelagic Fishery
```{r}
blue_fauna <- blue_water_fauna_and_fleets$fauna
blue_fleets <- blue_water_fauna_and_fleets$fleets
blue_resolution <- sqrt(blue_fauna[[1]]$patches)
```
We model our pelagic case study loosely on the characteristics of the Western
and Central Pacific Ocean (WCPO) tuna fisheries. Note that this is an illustrative example only and simulated stock status, species distributions, and projections presented here should not be interpreted as a indicative of the current or future state of the WCPO. We simulate trajectories of
`r n_distinct(names(blue_fauna))` species commonly caught in the region,
including both the highly abundant skipjack tuna and the heavily depleted
oceanic whitetip shark (*Carcharhinus longimanus*, Carcharhinidae) (@fig-blue-ssb). We use
publicly available data on catch-per-unit-effort of each of these species from
the WCPO as a very rough proxy for baseline habitat distributions, noting that
where possible, fishery-independent abundance indices would be preferable
(@fig-blue).
These pelagic species are caught by a longline fleet that primarily targets
large adult tunas such as bigeye and yellowfin (*Thunnus albacares*, Scombridae) for high-grade consumption, and a
purse-seine fishery that primarily targets skipjack tunas for bulk canning.
Contact selectivities were modeled as logistic for the longline fleet, and
dome-shaped for the purse-seine fleet. Both fleets operate under open-access dynamics with an effort
cap. The effort cap was set at the level of effort that resulted in the desired
levels of SSB/SSB0 for each species under open-access dynamics (@fig-blue),
intended to simulate a scenario where managers step in to prevent further
expansion of fishing effort in a fully developed fishery. For
forward-simulation, open-access dynamics can result in effort decreasing in
response to profitability, but cannot result in effort beyond the effort cap set
for each fleet.
For this exercise, we focused on using `marlin` to assess resilience of the
selected *Target Fishing* and *Rate* MPA design strategies to a climate-driven
range shift. Specifically, we simulate an extreme example where the centroid of
each population shifts northward at a rate of ~62km per year over a 20
year time horizon (@fig-blue-ssb). We designed MPA networks given the conditions
in the starting year, and then held that network constant over the years of the
experiment, running one simulation with and another without the climate-drive
range shift. We then compared the effects of this range shift on food production
and conservation outcomes from MPA networks designed based on the pre-range
shift world. We ran the pelagic simulation at a quarterly level ($\Delta_t$ =
1/4 year), and set the area of each cell to be roughly 97,000 KM^2^ across 144
patches each with a side length of roughly 311 KM, for a total area of
14e6 KM^2^, broadly commensurate with the area of the WCPO.
## Results
### Coastal Coral Reef Fishery
```{r}
#| label: coral-results
coral_mpa_experiments$mpas <- map(coral_mpa_experiments$results, "mpa")
coral_mpa_experiments$obj <- map(coral_mpa_experiments$results, "obj")
coral_results <- coral_mpa_experiments %>%
unnest(cols = obj)
coral_fleet_results <- coral_results %>%
pivot_longer(starts_with("fleet_"),
names_to = "fleet",
values_to = "fleet_yield") %>%
group_by(prop_mpa, fleet, placement_strategy) %>%
summarise(yield = sum(fleet_yield), biodiv = sum(unique(biodiv))) %>%
group_by(fleet, placement_strategy) %>%
mutate(delta_yield = yield / yield[prop_mpa == 0] - 1) |>
ungroup()
peak_coral_fleet_results <- coral_fleet_results |>
group_by(fleet) |>
filter(delta_yield == max(delta_yield)) |>
mutate(peak_yields = scales::percent(delta_yield))
tmp <- coral_results %>%
group_by(prop_mpa, placement_strategy) %>%
summarise(biodiv = sum(biodiv), yield = sum(yield)) %>%
group_by(placement_strategy) %>%
mutate(delta_yield = yield / yield[prop_mpa == 0] - 1,
delta_biodiv = biodiv / biodiv[prop_mpa == 0] - 1) %>%
ungroup() %>%
mutate(placement_strategy= fct_relabel(placement_strategy, titler))
```
MPAs were capable of producing a range of positive and negative outcomes for
food security and conservation in the coral reef case study depending on the
design strategy used and the size of MPA implemented. Both of the MPA design
strategies were capable of increasing fisheries yield for
Fleet Two, up to a value of
`r peak_coral_fleet_results$peak_yields[peak_coral_fleet_results$fleet == "fleet_two"]`,
even when MPAs covered more than 50% of the simulated area. However Fleet One only benefited from MPAs under the *Spawning Ground*
design strategy, with a maximum increase of
`r peak_coral_fleet_results$peak_yields[peak_coral_fleet_results$fleet == "fleet_one"]`;
the *Target Fishing* design strategy produced a roughly linear decrease in
fishing yields as a function of increasing MPA size (@fig-coral-results A).
MPAs were uniformly beneficial to the spawning
biomass of all species under the *Target Fishing* design strategy, with the most
rapid increases in spawning biomass for the deep snapper population. The
*Spawning Ground* strategy primarily benefited the shallow snapper population,
producing little change in the grouper population and decreasing spawning
biomass of both the deep snapper and reef shark populations for MPA sizes
covering less than 50% of the simulated area (@fig-coral-results B).
In total, the *Spawning Ground* strategy was capable or providing net increases