Derek Mitchell has another research article that addresses hive configuration and its impacts on humidity. Apparently, it also covers the implications of higher humidity on varroa reproduction.
https://royalsocietypublishing.org/doi/10.1098/rsif.2019.0048
From the abstract:
"It is highly likely that honeybees, in temperate climates and in their natural home, with much smaller thermal conductance and entrance, can achieve higher humidities more easily and more frequently than in man-made hives. As a consequence, it is possible that Varroa destructor, a parasite implicated in the spread of pathogenic viruses and colony collapse, which loses fecundity at absolute humidities of 4.3 kPa (approx. 30 gm−3) and above, is impacted by the more frequent occurrence of higher humidities in these low conductance, small entrance nests."
https://royalsocietypublishing.org/doi/10.1098/rsif.2019.0048
From the abstract:
"It is highly likely that honeybees, in temperate climates and in their natural home, with much smaller thermal conductance and entrance, can achieve higher humidities more easily and more frequently than in man-made hives. As a consequence, it is possible that Varroa destructor, a parasite implicated in the spread of pathogenic viruses and colony collapse, which loses fecundity at absolute humidities of 4.3 kPa (approx. 30 gm−3) and above, is impacted by the more frequent occurrence of higher humidities in these low conductance, small entrance nests."