Understanding how the world's biodiversity is organized and how it changesacross geographic regions is critical to predicting the effects of global change 1 . Ecologists have long documented that the world's terrestrial fauna is organized hierarchically in large regions -or realms -and continental scale subregions 2-6 , with boundaries shaped by geographic and climatic factors 2,7 . However, little is known about how global biodiversity is assembled below the continental level and the factors, including the potential role of human impacts, triggering faunistic differences as the biogeographical scale becomes smaller. Here we show that the hierarchical organization of global zoogeographical regions extends coherently below the region level to reach a local scale, and that multiple determinants act across varying spatial and temporal scales. Among these determinants, anthropogenic land use during the Late Holocene stands out showing a footprint across biogeographical scales and explaining 22% of the faunistic differences among the larger bioregions. The Late Holocene coincided with the development of large cities and substantial transformation of ecosystems into agricultural land 8,9 . Our results show that past human activity has played a role in the global organization of present-day animal assemblages, leaving a detectable signal that warns us about significant time-lag effects of human-mediated impacts on biodiversity. The questions of how the world's biodiversity is organized, and why large-scale patterns of taxonomic diversity change through natural geographic regions have attracted the attention of naturalists since the early 19 th century 2,10-15 . The answers to these questions are important to satisfy our curiosity about the natural world, but have also become 3 critical to forecast the future of biodiversity in the face of global change 1 . A key step in understanding the organization of biodiversity is the assemblage of regions based on their shared elements 14 . Alfred R. Wallace was among the first to propose that the world's fauna is organized hierarchically in broad regions shaped by geographic and climatic factors 2 . About 150 years later, the development of multivariate analytical techniques has led to the revaluation of Wallace's proposal 3-6 and refining of the extrinsic factors explaining the major dissimilarities among zooregions 7 . However, biogeographic boundaries and the signal of evolutionary processes associated to species isolation are not so evident at smaller scales, and importantly, still remain globally unexplored. Smaller regions, which are generally the units of conservation actions, contain more similar biota and thus, the factors determining faunistic dissimilarities among them are likely to be more diverse and include spatial and taxonomic idiosyncracies 7,16,17 .We hypothesize that global biodiversity patterns can be characterized by a hierarchical system of biogeographic regions extending from global to local scales, with regions at different levels explained by determinants...