Two eminent ecologists, the late Robert MacArthur of Princeton University and E. 0. Wilson of Harvard, developed a theory of "island biogeography" to explain such uneven distributions. They proposed that the number of species on any island reflects a balance between the rate at which new species colonize it and the rate at which populations of established species become extinct. If a new volcanic island were to rise out of the ocean off the coast of a mainland inhabited by 100 species of birds, some birds would begin to immigrate across the gap and establish populations on the empty, but habitable, island. The rate at which these immigrant species could become established, however, would inevitably decline, for each species that successfully invaded the island would diminish by one the pool of possible future invaders the same 100 species continue to live on the mainland, but those which have already become residents of the island can no longer be classed as potential invaders.