Arthropodization in the Hirudinea: evidence for a phylogenetic link with insects and other Uniramia?

The Hirudinea have a number of arthropod-like characters, including a true haemocoel and compound eyes. This ‘arthropodization’ may have bearing on the origin of the arthropod phylum Uniramia. Certain traits shared by Clitellata and Uniramia are interpreted as primitive and compatible with the view that a monophyletic link exists between these two groups: ‘internal’ fertilization, egg protected by cocoon/chorion, egg yolky with direct development not invoking a planktotrophic larval stage, formation and fate of presumptive areas in early embryology, uniramous lobopodial-like structures and the presence of uniramous mouthparts. Still other traits shared by Hirudinea and Hexapoda are interpreted as advanced and evidence for a paraphyletic relationship between leeches and pterygote insects: oogenesis with nurse cells, cephalization and segmental constancy. It is proposed that a major theme in uniramian evolution is the invasion of land from freshwater by a pre-lobopodial clitellate-like ancestor, the lobopodium being an adaptation to terrestrial locomotion. These views give new significance to the poorly studied fossil segmented worms from freshwater and terrestrial palaeohabitats.