The Lesser kestrel (Falco naumanni) evolved as a separate species in the Old-World kestrel radiation starting in the late Miocene. Therefore, this urban colonial raptor has only become a major town dweller recently in its evolutionary history, and just in part of its breeding range (the Western Palearctic). Today, more than 95% of lesser kestrel colonies in Spain and other Mediterranean countries are on buildings, and the few remaining are on rocky outcrops, that may have been the original nesting substrate for this cavity-nesting bird. Lesser kestrel fossils are well represented in cave sites, and their paleontological distribution, spanning to form the Early Paleolithic to the Epipaleolithic, agrees well with its current breeding distribution. According to classical sources, such as the works of Columella and Pliny the Elder, and the presence of a skeletal remain in a Roman villa near Madrid, lesser kestrels may have nested in buildings and in urban settings for at least 2000-2500 years. However, there are no surviving colonies in structures older than 1400 years in Andalusia, nor in Spain. For a sample of 349 colonies on ancient buildings, a majority of the structures had been erected in the 15 th and 16 th centuries, this putting a time limit of 400-500 years to the existence of those seemingly immemorial colonies. For specific towns and buildings, written references for the presence of lesser kestrel colonies do not go back more than 200 years. In fact, the Cathedral of Sevilla may be the structure with the longest continuous occupation by lesser kestrels recorded up to present time, from at least Cook's (1934) notes to the 2020 breeding season. Lesser kestrels were possibly too common in human settlements in the past as to be noted as special. This may explain the scarcity of references to the species until the 19th century. In any case, the same lack of concretion affects the other major Eurasian urban birds, as no timeline exist for the urbanization process of any other bird species. We propose that lesser kestrels became urban breeders when both adequate cavities and cereal agriculture were developed in their breeding range, several millennia ago. However, specific urban colonies, contrasting with the stability of geological substrates, have adaptively moved from building to building when older ones become ruinous or were rebuilt, and new options became available throughout History.