A red nucleus, defined by its relative position in the tegmentum mesencephali, its contralateral rubrospinal or rubrobulbar projections and by crossed cerebellar afferents, is found in terrestrial vertebrates and certain rays. A crossed rubrospinal tract occurs in anurans, limbed urodeles and reptiles, birds and mammals, but is apparently absent in boid snakes, caecilians and sharks. A distinct rubrospinal tract is found in certain rays which use their enlarged pectoral fins for locomotion. A crossed tegmentospinal tract, possibly a rubrospinal tract, is found in lungfishes. Although evidence was presented for a rubrospinal tract in more advanced snakes, the available experimental data in lower vertebrates suggest that the presence of a rubrospinal tract is related to the presence of limbs or limb-like structures. In the connectivity of the red nucleus in terrestrial vertebrates, 'levels' of complexity can be distinguished, paralleled by the development of the cerebellum. These 'grades of organization' are probably related to the type of motor performance the particular terrestrial vertebrates are capable of.