DOI: 10.5061/dryad.1g1jwsv3d
title: Toward greater realism in inclusive fitness models: the case of caste fate conflict in insect societies

description: In the field of social evolution, inclusive fitness theory has been
 successful in making a wide range of qualitative predictions on expected
 patterns of cooperation and conflict. Nevertheless, outside of sex ratio
 theory, inclusive fitness models that make accurate quantitative
 predictions remain relatively rare. Past models dealing with caste fate
 conflict in insect societies, for example, successfully predicted that if
 female larvae can control their own caste fate, an excess should opt to
 selfishly develop as queens. Available models, however, were unable to
 accurately predict levels of queen production observed in Melipona bees –
 a genus of stingless bees where caste is self-determined – as empirically
 observed levels of queen production are ca. two times lower than the
 theoretically predicted ones. Here, we show that this discrepancy can be
 resolved by explicitly deriving the colony-level cost of queen
 overproduction from a dynamic model of colony growth, requiring
 incorporation of parameters of colony growth and demography, such as the
 per-capita rate at which new brood cells are built and provisioned, the
 percentage of the queen’s eggs that are female, costs linked with worker
 reproduction and worker mortality. Our revised model predicts queen
 overproduction to more severely impact colony productivity, resulting in
 an evolutionarily stable strategy (ESS) that is ca. half that of the
 original model, and is shown to accurately predict actual levels of queen
 overproduction observed in different Melipona species. Altogether, this
 shows how inclusive fitness models can provide accurate quantitative
 predictions, provided that costs and benefits are modelled in sufficient
 detail and are measured precisely.
description: This zip directory contains all the necessary files to reproduce the
 findings reported in our associated paper, titled "Toward greater
 realism in inclusive fitness models: the case of caste fate conflict in
 insect societies". Citation: Ferreira, H. M., Alves, D. A., Cool, L.,
 Oi, C. A., Oliveira, R. C., & Wenseleers, T. (2024). Toward greater
 realism in inclusive fitness models: the case of caste fate conflict in
 insect societies. Evolution Letters, qrad068. The contents include: ### I.
 Raw data files used for the statistics: 1.
 1_Queen_male_production_raw_data.csv : Species names, Population (refers
 to each sampled location), Locality, Ntcombs (number of combs accessed),
 Ntcolony (total number of colonies accessed), Ncolony (Number of colonies
 accessed per row), Ncomb(number of combs accessed per row), Idcolony
 (individual colony ID (xx_pooled refers to the fact that is lacking
 individual data per colony/comb), Tmonths (total number of months in which
 the colonies/combs were accessed), month (specific month in which the
 colonies/combs were accessed), NCells (number of cells accessed), Nfemales
 (number of females), Nworkers (number of workers), Nmq (number of queens
 reared in worker cells), Nrq (Number of queens reared in royal cells),
 Nmales (number of males), ref (reference to the article/theses/abstracts
 where the data was collected from). NA refers to missing data. 2.
 2_cell_building.csv : Species (Species name), population (refers to each
 sampled location), sampling_locality (City and/or Country where the
 colonies were sampled), colony (Colony individual ID), colony_size (colony
 size measured by number of workers), reference_colony_size (literature
 used to obtain colony size), observation_days (observation period in
 days), worker_days (colony_size*observation days, used to calculate how
 many cells were built per day, per worker), total_cells_built (total
 number of cells built over the mentioned period), cells_built_per_day
 (number of cells built per day), cells_built_per_day_per_worker (number of
 cells built per day per worker), total_nr_study_colonies_per_species
 (total number of colonies accessed per species), reference_cell_building
 (literature from where it was obtained information regarding the cell
 building for the mentioned species), notes (notes on gathering and
 processing information from the literature). 3. 3_worker life
 expectancy.csv : species, population (refers to each sampled location),
 nstudies (number of sudies reporting worker life expectancy),
 avg_worker_lifespan (average worker life span in days), n (number of
 workers from which it was reported the worker life expectancy), ncols
 (number of colonies), reference & full reference (refers to the
 literature from where it was obtained the data on the worker life
 expectancy). 4. 4_worker-reproduction.csv : species, population (refers to
 each sampled location), prop_WPM (proportion of Worker Produced Males),
 Nwpm (number of worker produced males), Nm (number of males for which the
 parentage was dtermined), Na (effective number of males for which the
 parentage could be assigned given the power of the technique used), Cols
 (number of colonies), Method (method used to infer male parentage:
 allozyme markers (A), behavioural data (B), microsatellite markers (M)),
 References (citations for articles providing data on worker reproduction).
 ### II. R code & R environment 1. The script
 "caste_conflict_model_R_script_FINAL.r" was used to obtain the
 stats on the raw data. All the packages required are mentioned in the
 script that was developed on R version 4.1.1 (R Core Team 2021). 2. R
 environment file: "environment.RData". ### III. Mathematica
 Notebook 1. A notebook file "caste fate conflict model_FINAL.nb"
 for developing the model on caste conflict. This file is compatible with
 Wolframe Mathematica version 10.0. A description of all the parameters is
 given in the notebook file. 2. PDF version of the Mathematica notebook:
 "caste fate conflict model_FINAL.pdf" ### IV. All the remaining
 files were generated by running the R script on R version 4.1.1. For any
 query or further assistance, please contact: Helena Mendes Ferreira
 ( or
 ).