> The upward tilt of combs is likely a reflection of the fact that honey bees build their nests within enclosures, and not out in the elements

Wrong. Only some species of honeybee are cavity nesting and even those often nest in unprotected locations.

"The honeybees comprise three groups with respect to nesting: the dwarf honeybees, A. andreniformis and A. florea, are single comb, open-nesting species; the medium-sized species, A. cerana, A. koschevnikovi, A. mellifera, A. nigrocincta and A. nuluensis, are multiple comb cavity-nesting bees; and the giant honeybees, A. dorsata and A. laboriosa, are also single comb open-nesting species/"

The authors also state:

"The shape of the honeybee cell does not have its celebrated regularity; its economy is a teleological myth. The entire history of the honeybee cell in natural history, geometry and philosophy is the story of centuries-old misconceptions."

-- Hepburn, H. R., Pirk, C. W. W., & Duangphakdee, O. (2016). HONEYBEE NESTS. Springer-Verlag Berlin An.

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