Chimpanzees ( Pan troglodytes ) are, along with bonobos, humans' closest living relatives. The advent of diffusion tractography in recent years has allowed a resurgence of comparative neuroanatomical studies in humans and other primate species. Here, we offer, in comparative perspective, the first chimpanzee white matter atlas, coupled with surface projection maps of these major white matter tracts, constructed from in vivo chimpanzee diffusion-weighted scans. Comparative white matter atlases provide a useful tool for identifying neuroanatomical differences and similarities between humans and other primate species. Until now, comprehensive fascicular atlases have been created for humans ( Homo sapiens ), rhesus macaques ( Macaca mulatta) , and several other nonhuman primate species, but never in a nonhuman ape. Information on chimpanzee neuroanatomy is essential for understanding the anatomical specializations of white matter organization that are unique to the human lineage.We identify the major white matter fibers by means of standardized anatomical landmarks that can be directly compared to those of previous studies in the human and macaque monkey brain [8] . These landmarks are used to create 'recipes' for each tract in a standardized brain template, and as such, can be easily transformed into different datasets of individual scans. We use these recipes to reconstruct the major white matter tracts in 29 in vivo diffusion-weighted MRI datasets using probabilistic tractography [9] . The nature of these recipes also means that future modifications can easily be incorporated into the atlas, as can competing recipes to compare claims between rival definitions of any given tract. A specialized tool for this, compatible with our data, has recently been released [10] .Cortical regions can be characterized and compared across species by describing their unique connectivity profiles, or "connectivity fingerprints" [11,12] . Therefore, for a tract atlas to provide insight into grey matter organization, it is informative to characterize the relationship of each white matter tract to cortex. Using the reconstructed tracts, we created cortical projection maps on a standardized chimpanzee template. We also created corresponding maps for the better understood human and macaque monkey to allow for direct comparisons.We discuss some interesting similarities and differences in the organization of major fasciculi, and their patterns of cortical terminations. Further, we offer resources for neuroanatomists interested in the evolution of the human and chimpanzee brain: an atlas of 24 major white matter tracts in chimpanzees, with directly comparable tracts in humans and macaques, surface projection data from these tracts, and tractography "recipes" for reproducing these tracts. These resources are made available to the scientific community in online repositories (www.neuroecologylab.org).
Results
OverviewWe have created tractography recipes for 24 tracts in the chimpanzee brain as well as corresponding recipes for the human and ma...