bayesTLS is an R package for fitting joint Bayesian four-parameter logistic
(4PL) models to thermal death time (TDT) and thermal load sensitivity (TLS) data.
It works with raw counts or continuous proportions, extracts classical quantities
such as z, CTmax, Tcrit, and LT curves as posterior summaries, and provides
tools for uncertainty propagation, group contrasts, heat-injury prediction, and
bundled case-study data.
Please cite the companion paper when using bayesTLS:
Noble, D.W.A., Arnold, P.A., Nakagawa, S. & Pottier, P. (2026). A flexible modelling framework for estimating thermal tolerance and sensitivity. bioRxiv. https://doi.org/10.64898/2026.07.16.738378
This preprint is the primary citation for both the statistical framework and
the bayesTLS package. You can also run citation("bayesTLS") in R for the
formatted reference and a BibTeX entry.
The analytical workflow ships as an installable R package, available on CRAN. Install the released version with:
install.packages("bayesTLS")
library(bayesTLS)To install the development version from GitHub instead:
# install.packages("remotes") # if needed
remotes::install_github("daniel1noble/bayesTLS")The package depends on brms (which needs a
Stan backend; cmdstanr is recommended). All
other imported dependencies (dplyr, ggplot2, MASS, patchwork,
posterior, tibble) are CRAN packages and resolve automatically. The classical
two-stage beta-binomial Stage 1 additionally uses the suggested
glmmTMB package.
The coding workflow is decoupled — each step is a standalone function so users can stop, swap, or skip at any stage:
| Function | Role |
|---|---|
standardize_data() |
Rename user columns to standard names; supports count responses (survived/dead/total) and continuous proportions; records response type + metadata. |
make_4pl_priors() / make_4pl_formula() |
Disjoint-bounds reparameterised 4PL; asymptote bounds adjustable for sublethal/PSII ranges. |
fit_4pl() |
Joint Bayesian 4PL via brms. Family resolved from the response — beta_binomial(identity) for counts, Beta(identity) for proportions. Centred temperature; temp_effects selects which 4PL parameters depend on temperature (all four by default; "mid" for sparse designs). Two parameterisations: midpoint (default) and direct CTmax/z (supply ctmax =/z = formulas). To compare moderator groups (species, life stage, …) in one joint fit, use by = "<moderator>" (midpoint — fits ~ temp_c * moderator on all four sub-parameters) or ctmax = ~ 0 + <moderator> (direct); the moderator is recorded so downstream readers auto-group. mid = gives explicit control of the midpoint sub-model. |
tls() / tls_z() / tls_ctmax() / tls_tcrit() |
One call to derive z, CTmax, and (lethal data only, lethal = TRUE) Tby =; sex, life stage, clone, …) from any fitted 4PL — a bayes_tls workflow or a hand-written brms 4PL — all from one consistent posterior. z and CTmax share one posterior subsample; seed = makes the subsample + Ttarget_surv picks the relative (default) or absolute threshold. |
derive_z(), derive_tdt_curve(), derive_temperature_for_duration(), derive_tdt_landscape() |
Exported TDT primitives (the building blocks tls() composes, and available directly for advanced use such as the per-temperature local z(T) profile from derive_z()); all support per-group results via by =. |
extract_4pl_pars(), tdt_parameter_table(), tdt_quantile(), summarise_observed_survival() |
Per-draw 4PL parameters, natural-scale posterior parameter tables (group-aware via by =), TDT-friendly quantile summaries, and observed-survival cell summaries (mean ± SE per temp × duration). |
predict_survival_curves(), predict_heat_injury(), repair_rate_schoolfield() |
Posterior survival curves and heat-injury accumulation under a temperature trace (both per-group via by =), with an optional Sharpe–Schoolfield repair kernel supplied by repair_rate_schoolfield(). |
make_temperature_scenarios(), planted_dose_from_trace() |
Reference temperature traces and the analytical HI integral for validation. |
ts_stage1(), ts_stage2(), ts_ci(), ts_curve() |
The classical two-stage TDT pipeline for side-by-side comparison: per-temperature dose-response (stats::glm binomial or glmmTMB beta-binomial) → OLS of log10(LT50) on temperature → z, CTmax, T |
diagnose_tdt_fit(), bayes_R2_tls() |
Sampling diagnostics (R-hat, ESS, divergences) and Bayesian R^2 summaries for a fitted workflow. |
plot_survival_curves(), plot_tdt_curve(), plot_tdt_landscape(), plot_heat_injury(), plot_temperature_scenarios(), plot_temperature_density(), plot_repair_rate(), theme_tdt() |
Plotting helpers for survival curves, TDT curves, tolerance landscapes, heat injury, temperature traces/densities, and repair TPCs, with a shared project theme. Where a fitted curve/surface is overlaid on data (plot_survival_curves(), plot_tdt_landscape(), plot_tdt_curve()), it is by default capped at each group's longest observed exposure (shorter durations still shown) and, for grouped fits, faceted with free per-group axes (clip_to_observed, facet_scales); pass clip_to_observed = FALSE for the full prediction grid. |
get_brmsfit(), get_tls_est(), get_4pl_est(), get_surv_draws(), get_hi_draws(), has_fit(), … |
Accessors for the underlying fit, posterior draws, and posterior summaries. get_tls_est() pulls draws or summary for any/all TLS quantities from a tls() object, merged on .draw so quantities keep their joint pairing (for contrasts and bivariate plots); get_4pl_est() is its 4PL-shape counterpart (draws or summary of low/up/k/mid from a fitted workflow). |
clock_to_minutes(), format_interval() |
Utilities: parse clock-time strings to elapsed minutes; format a posterior median with its interval as median [lower, upper]. |
Full reference: ?fit_4pl, ?tls, ?predict_heat_injury, etc.
Four publicly available datasets are included with the bayesTLS package to make it easier to reproduce analyses and results in the paper and for testing purposes. These datasets are:
| Dataset | Help | Endpoint |
|---|---|---|
snowgum_psii |
?snowgum_psii |
Snowgum leaf PSII (continuous proportion) |
dsuzukii |
?dsuzukii |
Drosophila suzukii multi-trait TDT (lethal, knockdown, fertility; per individual) |
zebrafish_o2 |
?zebrafish_o2 |
Zebrafish lethal TDT across an oxygen gradient (survival counts) |
aphid_tdt |
?aphid_tdt |
Cereal-aphid lethal TDT — three species × ages, heat + cold branches (survival counts) |
Each dataset can be loaded easily using data(aphid_tdt) (as an example of loading the aphid_tdt dataset). If you want other datasets loaded then simply replace aphid_tdt. If you want to learn more about a dataset you can explore it's helpful ?aphid_tdt.
This repository also contains the code needed to reproduce the companion paper,
its supplement, and the simulation results. To do this, install the package from
this repository, render the Quarto documents through the Makefile, and use the
tests to verify the package functions used by the manuscript and supplement. All
code chunks are provided in ms/ms.qmd and ms/supplement.qmd; use make supp
to render the supplement and scripts/simulations/run_simulations.R to rerun the
full two-stage-bias simulation suite. The large cached artifacts the renders read
(simulation outputs and brms model fits) are not stored in the repository;
make data downloads them from the project's public OSF deposit (see
Fetching cached outputs below) so you need
not re-fit or re-simulate. Run all scenarios with
Rscript scripts/simulations/run_simulations.R, or a subset by passing scenario
labels, e.g. Rscript scripts/simulations/run_simulations.R scen9_tmax_060. Each
scenario and the per-simulation pipeline are visible in that one script; the
reusable building blocks live in scripts/simulations/sim_functions.R.
Use this map to find the files and outputs involved in reproduction:
Makefile— main entry point for rendering the manuscript and supplement.ms/ms.qmd— manuscript source file; renders to_output/ms.html,_output/ms.docx, and_output/ms.pdf.ms/supplement.qmd— supplement source file; renders to_output/supp.html,_output/supp.docx, and_output/supp.pdf.R/— package source for the analysis functions used by the paper; see Key functions.data/— bundled package datasets as.rdafiles.inst/extdata/— source CSV files used to build the bundled datasets.data-raw/make_datasets.R— script that rebuilds the.rdadatasets from the source CSV files.output/models/— cachedbrmsfits (.rds) created and reused during renders.scripts/simulations/— the two-stage-bias simulation:run_simulations.R(scenarios + runner) andsim_functions.R(reusable building blocks).scripts/— other support scripts (presentation figures, document cleanup).tests/testthat/— fast unit tests plus optionalbrmsintegration tests.bib/— bibliography and the Ecology Letters CSL file.notes/— dated development notes for derivations, literature checks, and simulations.man/— roxygen-generated package help files._output/— final rendered manuscript and supplement files; created by the render commands below.
-
Install the package from this repository so the renders use the same code you are reproducing:
# install.packages("devtools") # if needed devtools::install()
-
Fetch the cached artifacts the renders read (simulation outputs and
brmsmodel fits), which are not stored in the repository:make data # results + model fits (~85 MB) -- enough to render the paper make data-full # ALSO the 2.6 GB raw/draws tier (only to re-aggregate sims)
make datais safe to re-run: a tier already present locally is skipped. No credentials are needed (the OSF deposit is public). -
Render the manuscript, supplement, or both from the repository root:
make all # manuscript and supplement, all formats make ms # manuscript only: _output/ms.{html,docx,pdf} make supp # supplement only: _output/supp.{html,docx,pdf}
You can also render one format at a time, for example
make ms-pdformake supp-html.The first supplement render may take several minutes because it fits or checks cached models. Subsequent renders reuse the
.rdsfiles inoutput/models/and should be much faster. -
Inspect the rendered files in
_output/. Usemake cleanto remove only_output/, ormake build-cleanto also remove the out-of-tree Quarto build cache at~/Library/Caches/tls-render/.
The simulation outputs and brms model fits the manuscript reads are large and
are not committed to the repository. They live in the project's public OSF
deposit (node c6dxy) as tiered zip archives, fetched
with make data:
| Command | Downloads | Size | Use |
|---|---|---|---|
make data |
results.zip + models.zip |
~85 MB | render the paper (default) |
make data-full |
the above plus sim_full.zip |
+2.6 GB | re-aggregate the raw simulation |
make data is idempotent — a tier already present under output/ is skipped
(pass TIER=--force to re-download), each download is md5-verified against
data-raw/osf_manifest.json, and the archives unpack straight into output/.
No credentials are required. Maintainers refresh the deposit with make osf-push
(needs an OSF_PAT token); see scripts/osf_publish.R.
-
Run the package tests if you want to verify the analysis functions separately from the manuscript render:
devtools::test() Sys.setenv(RUN_BRMS_TESTS = "true") devtools::test()
The first command runs the fast unit tests. The second also runs the gated
brmsintegration tests, which fit small cached models and check parameter recovery.
The manuscript and supplement are independent Quarto documents. There is no
project-level _quarto.yml; each .qmd file carries its own YAML so the files
can be rendered separately. The Makefile renders outside the Dropbox-synced
project directory, then copies only the final HTML, DOCX, and PDF files back to
_output/. This avoids conflicted intermediate files while preserving cached
model fits in output/models/.
Within a document, use Quarto's @ references as normal (@fig-X, @tbl-X,
@eq-X, @sec-X). Between the manuscript and supplement, use plain text, such
as "Equation 7 of the manuscript" in the supplement or "Figure S2" in the
manuscript. The supplement render automatically labels figures, tables, and
equations with an "S" prefix.
See CLAUDE.md §8a for more detail on the render architecture,
cross-reference handling, and the brms cache.
The supplement works through the same workflow on simulated data and four empirical examples:
- Brown shrimp — lethal and sublethal endpoints.
- Zebrafish — lethal TDT across life stages, fit as separate per-stage 4PLs
and as a joint model with
life_stageas a covariate. - Snow gum leaf PSII — continuous-proportion response fit with a Beta likelihood.
- Drosophila suzukii — multi-trait TDT data, including mortality, knockdown, and fertility endpoints.
The supplement also contains the two-stage-bias simulation results and sensitivity sweeps used to check the modelling framework.