“…Indirect measures of wild colony density ( n = 19, Table 2 ) were made by taking samples of honey bees from either the worker progeny of queens mated in an area of interest (Arundel et al., 2014 ; Jaffé et al., 2010 ; Moritz et al., 2007 ) or, more commonly, from drones trapped at drone congregation areas (DCAs; Arundel et al., 2013 ; Hinson et al., 2015 ; Jaffé et al., 2010 ; Moritz et al., 2007 , 2008 , 2013 ; Utaipanon, Holmes, et al., 2019 ; Utaipanon, Holmes, et al., 2021 ; Utaipanon, Schaerf, et al., 2021 ). The number of colonies within a given radius of the mating apiary or DCA is then inferred by the number of unique genotypes present in the sample of worker progeny or trapped drones, respectively.…”