Pits of California. These records date as far back as 100,000 years ago in the Pleistocene era, during which the pigeon's range extended to several western states that were not a part of its modern range. The abundance of the species in these regions and during this time is unknown. Ecology and behavior The passenger pigeon was nomadic, constantly migrating in search of food, shelter, or nesting grounds. In his 1831 Ornithological Biography, American naturalist and artist John James Audubon described a migration he observed in 1813 as follows: These flocks were frequently described as being so dense that they blackened the sky and as having no sign of subdivisions. The flocks ranged from only above the ground in windy conditions to as high as . These migrating flocks were typically in narrow columns that twisted and undulated, and they were reported as being in nearly every conceivable shape. A skilled flyer, the passenger pigeon is estimated to have averaged during migration. It flew with quick, repeated flaps that increased the bird's velocity the closer the wings got to the body. It was equally as adept and quick at flying through a forest as through open space. A flock was also adept at following the lead of the pigeon in front of it, and flocks swerved together to avoid a predator. When landing, the pigeon flapped its wings repeatedly before raising them at the moment of landing. The pigeon was awkward when on the ground, and moved around with jerky, alert steps. The passenger pigeon was one of the most social of all land birds. Estimated to have numbered three to five billion at the height of its population, it may have been the most numerous bird on Earth; researcher Arlie W. Schorger believed that it accounted for between 25 and 40 percent of the total land bird population in the United States. The passenger pigeon's historic population is roughly the equivalent of the number of birds that overwinter in the United States every year in the early 21st century. Even within their range, the size of individual flocks could vary greatly. In November 1859, Henry David Thoreau, writing in Concord, Massachusetts, noted that "quite a little flock of [passenger] pigeons bred here last summer," while only seven years later, in 1866, one flock in southern Ontario was described as being wide and long, took 14 hours to pass, and