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In this vignette, we demonstrate the new features that
paleopop
adds to poems
for modeling
populations over paleo-ecological time scales using the example of the
steppe bison in Siberia. The steppe bison was once distributed
throughout the Northern Hemisphere. It became regionally extinct in
Siberia 9,500 years ago, and it is thought to have gone completely
extinct in Canada 500 years ago. We will model the range dynamics of the
steppe bison in Siberia from the Last Glacial Maximum (21,000 years ago)
until 9,000 years before present. Along the way, we will learn how
paleopop
handles regions, model templates, simulations, and
results.
Here we load poems
and paleopop
and set the
number of parallel cores for simulation and an output directory for
results.
In this vignette, we will do the following:
Create a PaleoRegion
Generate dispersal rates and carrying capacities
Create the model template
Sample parameter space with Latin Hypercube Sampling
Run the simulations
Examine the results
While poems
implements a static region object, in
paleopop
we use something different, because regions on
paleo time scales are never static. Oceans rise, continents collide, and
the contours of our spatially explicit landscape shift. In the case of
the steppe bison in Siberia, the sea level rises over time.
paleopop
includes a temporally dynamic raster stack of
Siberia we can use to build a PaleoRegion
object. The
raster resolution is 2 by 2.
library(raster)
#> Loading required package: sp
raster::plot(paleopop::siberia_raster[[1]], main = "Siberia raster (LGM)",
xlab = "Longitude (degrees)", ylab = "Latitude (degrees)",
colNA = "blue")
raster::plot(paleopop::siberia_raster[[1001]], main = "Siberia raster (9000 BP)",
xlab = "Longitude (degrees)", ylab = "Latitude (degrees)",
colNA = "blue")
We can use this RasterStack
object as a template to
create a PaleoRegion
object, which will set the indices of
occupiable cells at each time step, and mask cells that are not
occupiable at a time step (in the case of this example, because those
cells are now underwater.) When we plot the region_raster
in the PaleoRegion
object, we see the maximum extent of the
region, showing all cells that are occupiable at any time step. The
temporal_mask_raster
method generates a
RasterStack
that marks occupiable cells at each time step
with a 1.
This part of the workflow is identical to poems
: we
create Generator
objects to dynamically generate dispersal
between grid cells, initial abundance, and carrying capacity at each
point in space and time.
We cut down on computational time during the simulations by generating ahead of time a matrix that describes the potential dispersal rates between every pairwise combination of cells in the study region. Dispersal rates are calculated using a distance-based dispersal function as well as, optionally, a conductance/friction landscape that maps out barriers to dispersal, such as ice or mountains. (We will omit the friction landscape for this example.)
dispersal_gen <- DispersalGenerator$new(region = region,
dispersal_max_distance = 500, # km
distance_classes = seq(10, 500, 10), # divide into bins
distance_scale = 1000, # sets units to km
dispersal_friction = DispersalFriction$new(), # empty
inputs = c("dispersal_p", "dispersal_b"),
decimals = 3)
dispersal_gen$calculate_distance_data(use_longlat = TRUE) # pre-calculate matrix of distances between cells
#> The extent and CRS indicate this raster is a global lat/lon raster. This means that transitions going off of the East or West edges will 'wrap' to the opposite edge.
#> Global lat/lon rasters are not supported under new optimizations for 4 and 8 directions with custom transition functions. Falling back to old method.
#> Loading required package: igraph
#>
#> Attaching package: 'igraph'
#> The following object is masked from 'package:raster':
#>
#> union
#> The following objects are masked from 'package:stats':
#>
#> decompose, spectrum
#> The following object is masked from 'package:base':
#>
#> union
#> Loading required package: Matrix
#>
#> Attaching package: 'gdistance'
#> The following object is masked from 'package:igraph':
#>
#> normalize
#> The extent and CRS indicate this raster is a global lat/lon raster. This means that transitions going off of the East or West edges will 'wrap' to the opposite edge.
#> Global lat/lon rasters are not supported under new optimizations for 4 and 8 directions with custom transition functions. Falling back to old method.
test_dispersal <- dispersal_gen$generate(input_values =
list(dispersal_p = 0.5,
dispersal_b = 200))$dispersal_data
head(test_dispersal[[1]])
#> target_pop source_pop emigrant_row immigrant_row dispersal_rate
#> 1 2 1 1 1 0.076
#> 2 3 1 2 1 0.065
#> 3 4 1 3 1 0.056
#> 4 9 1 4 1 0.028
#> 5 10 1 5 1 0.028
#> 6 11 1 6 1 0.026
Here we will generate a dynamic landscape based on a raster stack of
habitat suitability values (from 0, totally unsuitable, to 1, ideally
suitable). paleopop
has a spatiotemporally explicit habitat
suitability RasterStack
for Siberia from 21,000 - 9,000
years BP.
raster::plot(bison_hs_raster[[1]], main = "Bison habitat suitability (LGM)",
xlab = "Longitude (degrees)", ylab = "Latitude (degrees)",
colNA = "blue")
However, PaleoPop simulations are not based on habitat suitability values. Rather, they operate on initial populations and on carrying capacities. Therefore, we need to enter these habitat suitability landscapes into a generator that can translate them into an initial abundance landscape and into a dynamic carrying capacity landscape.
To do this, we must create a poems::Generator
object.
Because every simulation has different requirements, we must set our own
inputs, outputs, and generator functions to convert the inputs into the
outputs. Here, we will very simply use a maximum density parameter and
multiply it by the habitat suitability.
# Convert the habitat suitability raster to matrix
bison_hs <- bison_hs_raster[region$region_indices]
# Create the generator
capacity_gen <- Generator$new(description = "Capacity generator",
example_hs = bison_hs,
inputs = c("density_max"),
outputs = c("initial_abundance", "carrying_capacity"))
# indicate that initial abundance and carrying capacity are required
capacity_gen$add_generative_requirements(list(initial_abundance = "function",
carrying_capacity = "function"))
# define custom generative functions
capacity_gen$add_function_template("carrying_capacity",
function_def = function(params) {
round(params$density_max*params$example_hs)
},
call_params = c("density_max", "example_hs"))
capacity_gen$add_function_template("initial_abundance",
function_def = function(params) {
params$carrying_capacity[,1]
},
call_params = c("carrying_capacity"))
# test run
test_capacity <- capacity_gen$generate(input_values = list(density_max = 2000))
raster::plot(region$raster_from_values(test_capacity$initial_abundance), main = "Initial abundance",
xlab = "Longitude (degrees)", ylab = "Latitude (degrees)",
colNA = "blue")
We create a model template for simulation using the
PaleoPopModel
object. Let’s check the required attributes
for the object.
model_template <- PaleoPopModel$new()
model_template$model_attributes # all possible attributes
#> [1] "coordinates" "random_seed" "time_steps"
#> [4] "years_per_step" "populations" "initial_abundance"
#> [7] "transition_rate" "standard_deviation" "compact_decomposition"
#> [10] "carrying_capacity" "density_dependence" "growth_rate_max"
#> [13] "dispersal_data" "dispersal_target_k" "harvest"
#> [16] "harvest_max" "harvest_g" "harvest_z"
#> [19] "harvest_max_n" "human_density" "abundance_threshold"
#> [22] "occupancy_threshold" "results_selection"
model_template$required_attributes # required attributes to run simulations
#> [1] "time_steps" "years_per_step" "populations"
#> [4] "initial_abundance" "transition_rate" "standard_deviation"
#> [7] "carrying_capacity" "harvest" "results_selection"
Initial abundance, carrying capacity, and dispersal data will be provided by the generators above. Here we will define an environmental correlation matrix so there can be spatial autocorrelation in the simulations.
# Distance-based environmental correlation (via a compacted Cholesky decomposition)
env_corr <- SpatialCorrelation$new(region = region, amplitude = 0.4, breadth = 500)
correlation <- env_corr$get_compact_decomposition(decimals = 2)
Normally we would have spatiotemporally dynamic data on human density
over time, which the prey- and predator-dependent harvest function
translates into numbers of bison harvested. The harvest function in
paleopop
is defined by harvest_z
, which
determines whether harvesting is a Type II or Type III functional
response, harvest_g
, a constant added to the denominator
that reduces harvest rates, and harvest_max_n
, a ceiling on
maximum prey density that can be harvested.
In the interest of simplicity for this example, we will create a matrix of constant low human density.
Now we have all the components we need to build the model template, less the model attributes that will be sampled via Latin Hypercube Sampling later.
model_template <- PaleoPopModel$new(
simulation_function = "paleopop_simulator", # this is the default; made it explicit for the example
region = region,
# coordinates: you could use XY coordinates to define the region instead
time_steps = 1001,
years_per_step = 12, # generational length for bison
populations = region$region_cells, # extracts number of population cells
# initial_abundance: generated
transition_rate = 1.0, # rate of transition between generations
# standard_deviation: sampled
compact_decomposition = correlation,
# carrying_capacity: generated
density_dependence = "logistic",
# growth_rate_max: sampled
harvest = TRUE,
# harvest_max: sampled
harvest_g = 0.4, # constant
# harvest_z: sampled
# harvest_max_n: sampled
human_density = human_density,
dispersal_target_k = 10, # minimum carrying capacity that attracts dispersers
# dispersal_data: generated
# abundance_threshold: sampled
# initial_n: sampled
occupancy_threshold = 1, # threshold: # of populations for the species to persist
random_seed = 321,
results_selection = c(
# ema (expected minimum abundance), extirpation, occupancy, human_density,
"abundance", "harvested"
)
)
Here the workflow is the same as in poems
: we sample
along a range of values so as to create a prior, and use these to
parameterize different simulations. Here, we will sample each range of
values uniformly so as to create an uninformative
prior, but the LHS generator object offers options for sampling along
different distributions to create an informative prior. For the
sake of this example, we will sample 10 different parameter
combinations, but it is advisable to run thousands in order to fully
sample the parameter space.
nsims <- 10 # adjust to run your own example if desired
lhs_generator <- LatinHypercubeSampler$new()
lhs_generator$set_uniform_parameter("standard_deviation", lower = 0, upper = sqrt(0.06))
lhs_generator$set_uniform_parameter("growth_rate_max", lower = log(1.31), upper = log(2.84))
lhs_generator$set_uniform_parameter("abundance_threshold", lower = 0, upper = 500, decimals = 0)
lhs_generator$set_uniform_parameter("harvest_max", lower = 0, upper = 0.35)
lhs_generator$set_uniform_parameter("harvest_z", lower = 1, upper = 2)
lhs_generator$set_uniform_parameter("density_max", lower = 500, upper = 3250) # alias for harvest_max_n
lhs_generator$set_uniform_parameter("dispersal_p", lower = 0.05, upper = 0.25) # for the dispersal generator
lhs_generator$set_uniform_parameter("dispersal_b", lower = 65, upper = 145) # for the dispersal generator
sample_data <- lhs_generator$generate_samples(number = nsims, random_seed = 123)
sample_data$sample <- c(1:nsims)
head(sample_data)
#> standard_deviation growth_rate_max abundance_threshold harvest_max harvest_z
#> 1 0.12844192 0.8633636 471 0.16758685 1.510286
#> 2 0.02171397 0.5930831 407 0.26785857 1.034352
#> 3 0.06815201 0.7415355 323 0.33290269 1.459999
#> 4 0.23228272 0.9579419 396 0.30130572 1.641069
#> 5 0.14845673 0.4207378 136 0.17998030 1.854928
#> 6 0.21182926 0.5269072 215 0.07769187 1.336949
#> density_max dispersal_p dispersal_b sample
#> 1 619.5955 0.08969914 96.14441 1
#> 2 1505.6085 0.11640746 138.50153 2
#> 3 2241.5265 0.09977226 112.63579 3
#> 4 1915.4510 0.24870600 131.40983 4
#> 5 3237.3751 0.16170967 116.23608 5
#> 6 1045.6603 0.05308405 97.72835 6
Here we build a simulation manager to manage the sample data,
generators, and model template. The simulation manager will use the
paleopop_simulator
function to run simulations.
sim_manager <- SimulationManager$new(sample_data = sample_data,
model_template = model_template,
generators = list(capacity_gen,
dispersal_gen),
parallel_cores = parallel_cores,
results_dir = output_dir)
sim_manager$results_filename_attributes <- c("sample", "results")
run_output <- sim_manager$run()
run_output$summary
#> [1] "10 of 10 sample models ran and saved results successfully"
The simulation log, run_output
, can be examined for
error messages and failure indices if any of the simulations fail. There
is also a detailed simulation log created in the output directory.
We can explore the results of the simulations using the convenient
wrapper of the PaleoPopResults
object. Each
PaleoPopResults
object can hold the result of one
simulation, and it can generate outputs that were not specified in the
results selection above, such as extirpation times for each
population.
results1 <- PaleoPopResults$new(
results = readRDS(paste0(output_dir, "/sample_1_results.RData")),
region = region, time_steps = 1001
)
head(results1$extirpation)
#> [1] 1 1 1 1 2 2
We can also use a PaleoPopResults
object as a kind of
template for the ResultsManager
, an object that can
calculate summary metrics for many results all at once. As with the
generators, we must define our own custom metrics functions, because
each in silico experiment calls for a different set of metrics
to summarize the outputs. We can use the metrics generated by the
PaleoPopResults
object as the basis for our custom summary
metrics.
results_model <- PaleoPopResults$new(region = region, time_steps = 1001, trend_interval = 1:10)
metrics_manager <- ResultsManager$new(simulation_manager = sim_manager,
simulation_results = results_model,
generators = NULL) # don't need generators
metrics_manager$summary_metrics <- c("abundance_trend", "extinction_time")
metrics_manager$summary_functions <- list()
metrics_manager$summary_functions$extinction_time <- function(simulation_results) {
extinction_time <- -12*(1001 - simulation_results$all$extirpation)-9001 # converts timestep to years BP
if (is.na(extinction_time)) {
extinction_time <- -9001 # if NA, then it survived to the end of the simulation
}
return(extinction_time)
}
metrics_manager$summary_functions$abundance_trend <- function(simulation_results) {
abundance_trend <- simulation_results$all$abundance_trend
return(abundance_trend) # this will use the trend interval specified above
}
gen_log <- metrics_manager$generate(results_dir = output_dir)
You can examine the log for the summary metrics calculation to browse error messages for failed calculations.
Now that we’ve calculated some summary metrics, based on outputs from
PaleoPopResults
like extinction time and abundance trend,
we can examine the simulation results. Here I show the simulated
extinction times.
metrics_manager$summary_metric_data$extinction_time
#> [1] -20977 -20821 -11509 -13813 -9001 -11521 -14053 -9001 -9001 -9001
Both of these summary metrics, abundance trend over the first ten time steps and extinction time, could be a suitable metric to converge toward a validation target.
From here, you may continue with the poems
workflow, as
explained in the poems
vignette,
creating a Validator
object for pattern-oriented modeling
to validate the results of the simulation.
These binaries (installable software) and packages are in development.
They may not be fully stable and should be used with caution. We make no claims about them.