The Late Cretaceous eutherian Zalambdalestes reveals unique axis and complex evolution of the mammalian neck

The typical mammalian neck consisting of seven cervical vertebrae (C1-C7) was established by the Late Permian in the cynodont forerunners of modern mammals. This structure is precisely adapted to facilitate movements of the head during feeding, locomotion, predator evasion, and social interactions. Eutheria, the clade including crown placentals, has a fossil record extending back more than 125 million years revealing significant morphological diversification in the Mesozoic. Yet very little is known concerning the early evolution of eutherian cervical morphology and its functional adaptations. A specimen of Zalambdalestes lechei from the Late Cretaceous of Mongolia boasts exceptional preservation of an almost complete series of cervical vertebrae (C2-C7) revealing a highly modified axis (C2). The significance of this cervical morphology is explored utilizing an integrated approach combining comparative anatomical examination across mammals, muscle reconstruction, geometric morphometrics and virtual range of motion analysis. We compared the shape of the axis in Zalambdalestes to a dataset of 88 mammalian species (monotremes, marsupials, and placentals) using three-dimensional landmark analysis. The results indicate that the unique axis morphology of Zalambdalestes has no close analog among living mammals. Virtual range of motion analysis of the neck strongly implies Zalambdalestes was capable of exerting very forceful head movements and had a high degree of ventral flexion for an animal its size. These findings reveal unexpected complexity in the early evolution of the eutherian cervical morphology and suggest a feeding behavior similar to insectivores specialized in vermivory and defensive behaviors in Zalambdalestes akin to modern spiniferous mammals.