>Ollie Visick seems to confirm that with his finding that 50% of feral colonies do not survive the winter and that the sites are then re-populated by swarms, probably from managed colonies (of which there are very large numbers in the area which he is studying).

This seems a reasonable critique - I would only clarify that Visick has not been focusing on the genetic basis of 'free living' colonies, rather simply on their potential quantity, nesting location and survival.

As noted in his most recent published work (https://link.springer.com/article/10.1007/s10841-023-00530-7):

'Our results show that TSIs provide valuable nesting habitat for wild-living honey bee colonies in southeast England. Based on an occupancy rate of 2% and a recent estimate of 2 million TSIs in England, we estimate that TSIs support 40,000 wild-living honey bee colonies across Britain and that wild-living colonies are a significant component of total colony numbers in areas with a high density of TSIs (> 50/km2).'

'Clearly, the colonies living in tree cavities in our study do not form an isolated wild population as all are within mating range of managed hives (up to 15 km, Jensen et al. 2005), but we still refer to honey bees living in tree cavities as “wild” at the colony-level. In some areas, colonies are not able to survive in the wild and many die during the winter and are replaced by swarms from managed hives the following spring and summer (Kohl et al. 2022). However, this is not the case in our study area. Most of the colonies in our study area are being monitored as part of a long-term study to determine colony survival. Preliminary data from the first three field seasons suggests that approximately half of all colonies in tree cavities survive the winter (ODV, unpublished data).'

He also refers to the 2012 Thompson paper in making inferences about free-living populations across Britain: 

'Our results show that approximately 2% of trees of special interest (TSIs), listed on the Ancient Tree Inventory (ATI) in southeast England, are used as nest sites by wild colonies. Based on this occupancy rate and the current number of TSI records, we estimate that there are approximately 350 wild colonies nesting in listed TSIs in our study area and approximately 3,300 across Britain. However, the ATI is far from complete and Nolan et al. (2022) estimate that there could be 2 million ancient and veteran trees in England alone (62% of Britain), so the number of wild colonies nesting in old trees in Britain could be over 40,000. This is a rough estimate based on the assumption that the results from our study area apply more widely in Britain. This is a reasonable assumption given that wild colonies are present throughout Britain (Thompson 2012; Seeley and Chilcott 2020) and that tree diversity in our study area largely reflects that of the whole ATI (Nolan et al. 2020).'

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