Which preposition to use with copulate

in Occurrences 1%

He tells us that boys burned bones and filth of various kinds to make a foul smoke, and that the smoke drove away certain noxious dragons which at this time, excited by the summer heat, copulated in the air and poisoned the wells and rivers by dropping their seed into them; and he explains the custom of trundling a wheel to mean that the sun, having now reached the highest point in the ecliptic, begins thenceforward to descend.

Which preposition to use with  copulate