bboutools is an R package to estimate Boreal Caribou recruitment, survival and population growth. Functions are provided to fit Bayesian or Maximum Likelihood (ML) models and generate and plot predictions.

Under the hood, the Nimble R package is used to fit heirarchical Bayesian and Maximum Likelihood models. Model templates in Nimble use BUGS-like syntax.

Several anonymized data sets are provided in a separate R package, bboudata.

In order to use bboutools you need to install R (see below) or use the Shiny app.

Philosophy

bboutools is intended to be used in conjunction with tidyverse packages such as readr, dplyr to manipulate data and ggplot2 (Wickham 2016) to plot data. As such, it endeavors to fulfill the tidyverse manifesto.

Installation

In order to install R (R Core Team 2023) the appropriate binary for the users operating system should be downloaded from CRAN and then installed.

Once R is installed, the bboutools package can be installed from GitHub by executing the following code at the R console

install.packages("remotes")
remotes::install_github("poissonconsulting/bboutools")

The bboutools package can then be loaded into the current session using

Getting Help

To get additional information on a particular function just type ? followed by the name of the function at the R console. For example ?bb_fit_recruitment brings up the R documentation for the bboutools recruitment model fit function.

For more information on using R the reader is referred to R for Data Science (Wickham and Grolemund 2016).

If you discover a bug in bboutools please file an issue with a reprex (reproducible example) at https://github.com/poissonconsulting/bboutools/issues.

Providing Data

Once the bboutools package has been loaded, the next task is to provide some data. An easy way to do this is to create a comma separated file (.csv) with spreadsheet software. Recruitment and survival data should be formatted as in the following anonymized datasets and can be checked to confirm it is in the correct format by using the bboudata functions:

# Recruitment data
bboudata::bbd_chk_data_recruitment(bboudata::bbourecruit_a)
bboudata::bbourecruit_a
#> # A tibble: 696 × 9
#>    PopulationName  Year Month   Day  Cows Bulls UnknownAdults Yearlings Calves
#>    <chr>          <int> <int> <int> <int> <int>         <int>     <int>  <int>
#>  1 A               1990     3     9     1     1             0         0      0
#>  2 A               1990     3     9     5     1             0         0      0
#>  3 A               1990     3     9     4     1             0         0      0
#>  4 A               1990     3     9     2     0             0         0      0
#>  5 A               1990     3     9     6     0             0         0      0
#>  6 A               1990     3     9     4     1             0         0      0
#>  7 A               1990     3     9     5     0             0         0      0
#>  8 A               1990     3     9     2     0             0         0      0
#>  9 A               1990     3     9     3     2             0         0      1
#> 10 A               1990     3     9     4     0             0         0      1
#> # ℹ 686 more rows
# Survival data
bboudata::bbd_chk_data_survival(bboudata::bbousurv_a)
bboudata::bbousurv_a
#> # A tibble: 364 × 6
#>    PopulationName  Year Month StartTotal MortalitiesCertain MortalitiesUncertain
#>    <chr>          <int> <int>      <int>              <int>                <int>
#>  1 A               1986     1          0                  0                    0
#>  2 A               1986     2          8                  0                    0
#>  3 A               1986     3          8                  0                    0
#>  4 A               1986     4          8                  0                    0
#>  5 A               1986     5          8                  0                    0
#>  6 A               1986     6          8                  0                    0
#>  7 A               1986     7          8                  0                    0
#>  8 A               1986     8          8                  0                    0
#>  9 A               1986     9          8                  0                    0
#> 10 A               1986    10          8                  0                    0
#> # ℹ 354 more rows

All columns should be included and column names should not be changed.

The .csv file can then be read into R using the following

data <- read_csv(file = "path/to/file.csv")

Recruitment

The annual recruitment in boreal caribou population is typically estimated from annual calf:cow ratios.

bboutools fits a Binomial recruitment model to the annual counts of calves, cows, yearlings, unknown adults and potentially, bulls.

It is up to the user to ensure that the data are from surveys that were conducted at the same time of year, when calf survival is expected to be similar to adult survival.

Fit a Bayesian model

The function bb_fit_recruitment() fits a Bayesian recruitment model.

The start month of the biological year (i.e., ‘caribou year’) can be set with the year_start argument. By default, the start month is April. Data are aggregated by biological year (not calendar year) prior to model fitting.

The adult female proportion can either be fixed or estimated from counts of cows and bulls (i.e., Cows ~ Binomial(adult_female_proportion, Cows + Bulls)).

If the user provides a value to adult_female_proportion, it is fixed. The default value is 0.65, which accounts for higher mortality of males (Smith 2004). If adult_female_proportion = NULL, the adult female proportion is estimated from the data (i.e., Cows ~ Binomial(adult_female_proportion, Cows + Bulls)). By default, a biologically informative prior of Beta(65,35) is used. This corresponds to an expected value of 65%.

The yearling and calf female proportion can be set with sex_ratio. The default value is 0.5.

The model can be fit with random effect of year, fixed effect of year and/or continuous effect of year (i.e., year trend). The min_random_year argument dictates the minimum number of years in the dataset required to fit a random year effect; otherwise a fixed year effect is fit. It is not recommended to fit a random year effect with fewer than 5 years (Kery and Schaub 2011). A continuous fixed effect of year can be fit with year_trend = TRUE.

The user can set quiet = FALSE argument to see messages and sampling progress.

recruitment <- bb_fit_recruitment(bboudata::bbourecruit_a, year_start = 4, year_trend = TRUE, quiet = TRUE)

Convergence

Model convergence can be checked with the glance() function.

glance(recruitment)
#> # A tibble: 1 × 8
#>       n     K nchains niters nthin   ess  rhat converged
#>   <int> <int>   <int>  <int> <dbl> <int> <dbl> <lgl>    
#> 1    27     4       3   1000    10   993  1.00 TRUE

Model convergence provides an indication of whether the parameter estimates are reliable. Convergence is considered successful by default if rhat < 1.05. The rhat threshold can be adjusted by the user. rhat evaluates whether the chains agree on the same values. As the total variance of all the chains shrinks to the average variance within chains, r-hat approaches 1.
Output of glance() also includes ess (Effective Sample Size), which represents the length of a chain (i.e., number of iterations) if each sample was independent of the one before it. By default, thebboutoolsBayesian method saves 1,000 MCMC samples from each of three chains (after discarding the first halves). The number of samples saved can be adjusted with thenitersargument. Withnitersset, the user can simply increment the thinning rate as required to achieve convergence (i.e., by decreasingrhat`).

Summary

Various generic functions in bboutools can be used to summarize or interrogate the output of model fitting functions.

  • coef() and tidy() provide a tidy table of the coefficient estimates.
  • estimates() provides a list of the coefficient estimates.
  • augment() provides the data used.
  • model_code() provides the model code in BUGS-like syntax.
  • plot() provides traceplots for individual parameters.

The user can exclude individual random effect estimates from coefficient output.

tidy(recruitment, include_random_effects = FALSE)
#> # A tibble: 3 × 4
#>   term    estimate  lower   upper
#>   <term>     <dbl>  <dbl>   <dbl>
#> 1 b0       -1.46   -1.63  -1.29  
#> 2 bYear    -0.0979 -0.258  0.0696
#> 3 sAnnual   0.326   0.188  0.517

Keep in mind that any reference to ‘Year’ or ‘Annual’ in these summary outputs represent the caribou year, which can be set by the user within the fitting functions.

Priors

In general, weakly informative priors are used by default (Gelman, Simpson, and Betancourt 2017; McElreath 2016). The default prior distribution parameter values can be accessed from bb_priors_recruitment() and bb_priors_survival(). See the priors vignette for more information.

bb_priors_recruitment()
#>                         b0_mu                         b0_sd 
#>                            -1                             5 
#>                      bYear_mu                      bYear_sd 
#>                             0                             2 
#>                    bAnnual_sd                  sAnnual_rate 
#>                             5                             1 
#> adult_female_proportion_alpha  adult_female_proportion_beta 
#>                            65                            35

The default prior distribution for adult_female_proportion is Beta(65, 35) and the default prior distribution for the intercept (b0) is Normal(-1.5, 1), which is on the log scale. The user can change the priors by providing a named vector to the priors argument in the model fitting functions. The names must match one of the names in bb_priors_recruitment().

For example, less informative priors for adult_female_proportion (e.g., Beta(1, 1)) and b0 (e.g., Normal(0, 5)) can be supplied as follows.

recruitment <- bb_fit_recruitment(bboudata::bbourecruit_a, priors = c(adult_female_proportion_alpha = 1, adult_female_proportion_beta = 1, b0_mu = 0, b0_sd = 5))

If the user is interested in fitting models without any prior information, see bb_fit_recruitment_ml() and bb_fit_survival_ml(), which use a Maximum Likelihood approach (see more details below).

Survival

The annual survival in boreal caribou population is typically estimated from the monthly fates of collared adult females. bboutools fits a Binomial monthly survival model to the number of collared females and mortalities. The user can choose whether to include individuals with uncertain fates with the certain mortalities.

The function bb_fit_survival() fits a Bayesian survival model.

The survival model is always fit with a random intercept for each month. Otherwise, the year_start, year_trend, and min_random_year arguments have the same behaviour as bb_fit_recruitment() above.

If include_uncertain_mortalities = TRUE, the total mortalities is the sum of the certain mortalities and uncertain mortalities (‘MortalitiesCertain’ and ‘MortalitiesUncertain’ columns); otherwise, only certain mortalities are used to fit the model.

survival <- bb_fit_survival(bboudata::bbousurv_a, year_start = 4, quiet = TRUE)
tidy(survival, include_random_effects = FALSE)
#> # A tibble: 3 × 4
#>   term    estimate  lower upper
#>   <term>     <dbl>  <dbl> <dbl>
#> 1 b0         4.46  4.16   4.8  
#> 2 sAnnual    0.327 0.0325 0.675
#> 3 sMonth     0.278 0.0608 0.669

Predictions

A user can generate and plot predictions of recruitment, survival and population growth.

Recruitment is the adjusted recruitment using methods from (DeCesare et al. 2012). See the ‘analytical methods’ article for details.

Predictions of calf-cow ratio can also be made using bb_predict_calf_cow_ratio().

The sex ratio can be adjusted with sex_ratio.

Recruitment by year

predict_recruitment <- bb_predict_recruitment(recruitment, year = TRUE)
bb_plot_year_recruitment(predict_recruitment)

Recruitment for a ‘typical’ year

predict_recruitment_1 <- bb_predict_recruitment(recruitment, year = FALSE)
predict_recruitment_1
#> # A tibble: 1 × 6
#>   PopulationName CaribouYear Month estimate  lower  upper
#>   <chr>                <int> <int>    <dbl>  <dbl>  <dbl>
#> 1 A                       NA    NA   0.0862 0.0758 0.0976

Recruitment trend

predict_recruitment_trend <- bb_predict_recruitment_trend(recruitment)
bb_plot_year_trend_recruitment(predict_recruitment_trend)

Survival by year for a ‘typical’ month

predict_survival <- bb_predict_survival(survival, year = TRUE, month = FALSE)
bb_plot_year_survival(predict_survival)

Survival by month for a ‘typical’ year

The estimates show annual survival, i.e., if that month lasted the duration of the year.

predict_survival_month <- bb_predict_survival(survival, year = FALSE, month = TRUE)
bb_plot_month_survival(predict_survival_month)

Population Growth

A user can predict population growth (lambda) with bb_predict_growth(). The survival and recruitment models fit in the previous steps are used as input. It is important to ensure that survival and recruitment outputs share the same biological year (i.e., ‘caribou year’), which is set with the year_start argument in bb_fir_survival() and bb_fit_recruitment(). Details on how lambda is calculated can be found in the analytical methods vignette.

predict_lambda <- bb_predict_growth(survival = survival, recruitment = recruitment)

bb_plot_year_growth(predict_lambda) +
  ggplot2::scale_y_continuous(labels = scales::percent)
#> Scale for y is already present.
#> Adding another scale for y, which will replace the existing scale.

Population change (%) is calculated with uncertainty as the cumulative product of population growth.

predict_change <- bb_predict_population_change(survival = survival, recruitment = recruitment)
bb_plot_year_population_change(predict_change)

Maximum Likelihood

Maximum Likelihood (ML) models can be fit using the bb_fit_recruitment_ml() and bb_fit_survival_ml() functions. These functions take a few seconds to execute because Nimble must compile the model into C++ code. See the Nimble documentation for more information and comparison to TMB. Similar to Bayesian model fits, generic functions (e.g., tidy(), glance() and augment()) work on ML fit objects (class ‘bboufit_ml’).

recruitment_ml <- bb_fit_recruitment_ml(bboudata::bbourecruit_a, year_start = 4, year_trend = TRUE, quiet = TRUE)
glance(recruitment_ml)
#> # A tibble: 1 × 4
#>       n     K loglik converged
#>   <int> <int>  <dbl> <lgl>    
#> 1    27     2  -90.1 TRUE
survival_ml <- bb_fit_survival_ml(bboudata::bbousurv_a, year_start = 4, quiet = TRUE)

The ML estimates are comparable to estimates derived from the equivalent Bayesian models above. In general, ML models can be interpreted as Bayesian models with uninformative (e.g., uniform) priors (McElreath 2016).

tidy(recruitment_ml, include_random_effects = FALSE)
#> # A tibble: 2 × 4
#>   term  estimate  lower   upper
#>   <chr>    <dbl>  <dbl>   <dbl>
#> 1 b0      -1.43  -1.53  -1.33  
#> 2 bYear   -0.112 -0.211 -0.0128
tidy(survival_ml, include_random_effects = FALSE)
#> # A tibble: 3 × 4
#>   term    estimate  lower upper
#>   <chr>      <dbl>  <dbl> <dbl>
#> 1 b0         4.45  4.16   4.75 
#> 2 sAnnual    0.335 0.102  1.1  
#> 3 sMonth     0.24  0.0728 0.789

There is functionality in bboutools to generate predictions (i.e., derived parameters) from ML models. However, there is no functionality to get confidence intervals on predictions. This is a more straightforward task with Bayesian models.

bb_predict_survival(survival_ml)
#> # A tibble: 32 × 6
#>    PopulationName CaribouYear Month estimate lower upper
#>    <chr>                <int> <int>    <dbl> <dbl> <dbl>
#>  1 A                     1985    NA    0.873    NA    NA
#>  2 A                     1986    NA    0.883    NA    NA
#>  3 A                     1987    NA    0.83     NA    NA
#>  4 A                     1988    NA    0.89     NA    NA
#>  5 A                     1989    NA    0.875    NA    NA
#>  6 A                     1990    NA    0.871    NA    NA
#>  7 A                     1991    NA    0.878    NA    NA
#>  8 A                     1992    NA    0.891    NA    NA
#>  9 A                     1993    NA    0.868    NA    NA
#> 10 A                     1994    NA    0.842    NA    NA
#> # ℹ 22 more rows
growth <- bb_predict_growth(survival_ml, recruitment_ml)
bb_plot_year_growth(growth)

Note that ML models can struggle to converge when there are sparse data, especially with a fixed year effect. If these issues arise, the user can try estimating year as a random effect, continuous effect (year_trend), or fitting a Bayesian model.

Another possible source of convergence issues is initial values. By default, bboutools sets initial values based on the default priors used for parameters in the Bayesian models. The user can replace initial values for parameters using inits.

inits_ml <- bb_fit_recruitment_ml(bboudata::bbourecruit_a, inits = c(b0 = 1, sAnnual = 0.3))

Understanding bboufit objects

The bb_fit_survival() and bb_fit_recruitment() functions use a Bayesian approach and return objects that inherit from class bboufit.

Objects of class bboufit have four elements:

  1. model - the compiled Nimble model as created by nimble::nimbleModel().
  2. model_code - the model code in text format.
  3. samples - the MCMC samples generated fromnimble::runMCMC() converted to an object of class mcmcr.
  4. data - the survival or recruitment data provided.

These are the raw materials for any further exploration or analysis. For example, view trace and density plots with plot(fit$samples).

See mcmcr and mcmcderive for working with mcmcr objects, or convert samples to an object of class mcmc.list, e,g, with coda::as.mcmc.list for working with the coda R package.

The bb_fit_survival_ml() and bb_fit_recruitment_ml() functions use a Maximum Likelihood approach and return objects that inherit from class bboufit_ml.

Objects of class bboufit_ml have five elements:

  1. model - the Nimble model as created by nimble::nimbleModel().
  2. model_code - the model code in text format.
  3. mle - the Maximum Likelihood output as created by model$findMLE().
  4. summary - the summary of the Maximum Likelihood output as created by model$summary(mle).
  5. data - the survival or recruitment data provided.

See nimble for how to work with nimble model objects and Maximum Likelihood output.

References

DeCesare, Nicholas J., Mark Hebblewhite, Mark Bradley, Kirby G. Smith, David Hervieux, and Lalenia Neufeld. 2012. “Estimating Ungulate Recruitment and Growth Rates Using Age Ratios.” The Journal of Wildlife Management 76 (1): 144–53. https://doi.org/10.1002/jwmg.244.
Gelman, Andrew, Daniel Simpson, and Michael Betancourt. 2017. “The Prior Can Often Only Be Understood in the Context of the Likelihood.” Entropy 19 (10). https://doi.org/10.3390/e19100555.
Kery, Marc, and Michael Schaub. 2011. Bayesian Population Analysis Using WinBUGS : A Hierarchical Perspective. Boston: Academic Press. http://www.vogelwarte.ch/bpa.html.
McElreath, Richard. 2016. Statistical Rethinking: A Bayesian Course with Examples in R and Stan. Chapman & Hall/CRC Texts in Statistical Science Series 122. Boca Raton: CRC Press/Taylor & Francis Group.
R Core Team. 2023. R: A Language and Environment for Statistical Computing. Vienna, Austria: R Foundation for Statistical Computing. https://www.R-project.org/.
Smith, Kirby Gordon. 2004. “Woodland Caribou Demography and Persistence Relative to Landscape Change in West-Central Alberta.” 125.
Wickham, Hadley. 2016. ggplot2: Elegant Graphics for Data Analysis. Springer-Verlag New York. https://ggplot2.tidyverse.org.
Wickham, Hadley, and Garrett Grolemund. 2016. R for Data Science: Import, Tidy, Transform, Visualize, and Model Data. First edition. Sebastopol, CA: O’Reilly. https://r4ds.had.co.nz.