would be completely destroyed during digestion, which therefore hindered dispersal of seeds this way. Instead, passenger pigeons may have spread seeds by regurgitation, or after dying. Reproduction Other than finding roosting sites, the migrations of the passenger pigeon were connected with finding places appropriate for this communally breeding bird to nest and raise its young. It is not certain how many times a year the birds bred; once seems most likely, but some accounts suggest more. The nesting period lasted around four to six weeks. The flock arrived at a nesting ground around March in southern latitudes, and some time later in more northern areas. The pigeon had no site fidelity, often choosing to nest in a different location each year. The formation of a nesting colony did not necessarily take place until several months after the pigeons arrived on their breeding grounds, typically during late March, April, or May. The colonies, which were known as "cities", were immense, ranging from to thousands of hectares in size, and were often long and narrow in shape (L-shaped), with a few areas untouched for unknown reasons. Due to the topography, they were rarely continuous. Since no accurate data was recorded, it is not possible to give more than estimates on the size and population of these nesting areas, but most accounts mention colonies containing millions of birds. The largest nesting area ever recorded was in central Wisconsin in 1871; it was reported as covering , with the number of birds nesting there estimated to be around 136,000,000. As well as these "cities", there were regular reports of much smaller flocks or even individual pairs setting up a nesting site. The birds do not seem to have formed as vast breeding colonies at the periphery of their range. Courtship took place at the nesting colony. Unlike other pigeons, courtship took place on a branch or perch. The male, with a flourish of the wings, made a "keck" call while near a female. The male then gripped tightly to the branch and vigorously flapped his wings up and down. When the male was close to the female, he then pressed against her on the perch with his head held high and pointing at her. If receptive, the female pressed back against the male. When ready to mate, the pair preened each other. This was followed by the birds billing, in which the female inserted