Hubrecht, Macfarlane, Jensen and Willmer: on the nature and testability of four versions of the nemertean theory of vertebrate origins

Abstract

Four authors (A. A. W. Hubrecht, J. M. Macfarlane, D. D. Jensen and E. N. Willmer) have argued that nemerteans are the proximal invertebrate ancestors of vertebrates. While all these authors accept certain homologies (i.e. vertebrate pituitary and notochord with nemertean proboscis invagination and proboscis sheath or mesoderm), there is considerable variation among them in the derivation of the vertebrate nervous system, sense organs and foregut apertures. For example, Hubrecht suggests that the vertebrate nervous system developed from a solid dorsal nerve found in some nemerteans; Macfarlane suggests it developed by fusion of the left and right solid brains of nemerteans; Jensen suggests that the vertebrate tubular nervous system developed by the backward extension of the nemertean frontal organ and fusion with the nemertean dorsal and ventral brain ganglia. Modern EM, histochemical and biochemical methods can test empirically the four theories. Research testing these theories will also test the more popular acraniate theory, since that theory implies that vertebrate head structures evolved de novo in chordates and that no homologies should exist between vertebrate and nemertean head structures.