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The traditional view that tarsiers are more closely related to middle Tertiary European microchoerines is supported by a new examination of the gross anatomy of the basicranium. Tarsius shares with Necrolemur and Microchoerus a suite of characters spanning several subsystems of the skull: the craniofacial junction, bony proportions of the basicranium, morphology of the pterygoids and the temporomandibular joint. Comparisons with other omomyids, adapids, plesiadapiforms and modern primates suggest that the states and combinations common to Tarsius, Necrolemur and Microchoerus are derived. Fewer components can be established in Tetonius, Pseudoloris and Nannopithex, whereas only primitive tarsiiform traits are evident in Rooneyia. Functional interpretations lend confidence to the inference that these phenetically very similar traits are homologous in Tarsius and its ‘necrolemur’ allies, and demonstrate their coordination with other synapomorphies of the cranium and postcranium. Necrolemurs appear to have undergone an adaptive transition to a crepuscular life-style, relying on insects significantly, using a powerful incisal killing bite, employing specialized auditory and optical prey detection techniques, and stealthy sit-and-wait foraging habits involving an important degree of leaping locomotion. These behaviors would have been preadaptive to the biology of tarsiers, and are built upon a homologous framework of derived tarsiiform characteristics.