in the mentioned study they say

"There is scarce scientific information on this practice, but different methods have been tested for long-term mass winter storage of queens."

Of course, this depends on what you mean by "scarce." I think there is ample information, but the investigators evidently didn't find the important stuff.

> Woyke, et al., (1956) found [60] years ago that worker bees injured the queens when several of them were stored in a colony. The workers bite claws, tarsi, antennae and even wings. According to Jasinski (1986, 1987) 60 % of 354 queens stored for 1 week in queenless colonies were injured. He detected 26 different types of injuries. The first injuries concerned the arolium also called the food pad. 

Woyke, J. (1988). "Problems with queen banks."  American Bee Journal. April, 1988. 276-278.

comment: I was told of this by Dennis van Englesdorp about 20 years ago when we were both working at the Dyce Lab for Honey Bee Studies at Cornell University, so I assumed it was common knowledge among informed beekeepers.

More recently:

> Drones were stored in mailing cages in their own colonies and stranger colonies. The number of injuries and the death rate were checked twice, after 3 and 7 days of storage. In total, over 4,608 drones were examined. Nine different types of injuries were observed for the drones, with leg injuries being the most common – lack of segments of tarsus (ca. 70-75% of all injuries). Other types of injuries included black arolia, missing arolia, wing and antenna injuries. 

Zajdel, B. et al. (2020). Are Drones Injured during Storage in Own and Stranger Queenright Colonies (Apis mellifera carnica)?. Journal of Agricultural Science and Technology, 22(2), 453-463.

Peter L Borst
Ithaca, NY  USA

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