Shani and Pnina’s paper was published in PLoS Computational Biology!
https://journals.plos.org/ploscompbiol/article?id=10.1371/journal.pcbi.1007653
This methodological study was actually a pilot for our GWAS of cuticular hydrocarbons. We wanted to optimize our genomic sequencing approach, in order to maximize the statistical power of GWAS for a colony-level trait. The results from this study can inform any GWAS in any social insect species of any colony-level trait. That is: a trait that is expressed at the level of the colony rather than the individual. E.g. social organization, foraging behavior, hygienic behavior, etc.
The figure below shows the expected power of GWAS depending on choices of methodology – check out the paper for details!