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Fig. 9. Evolution of O/P specification mechanisms in clitellate annelids deduced from phylogenetic analysis. The relationship of the three Helobdella laboratory populations used here is based on cytochrome oxidase I sequence analysis by A. Bely (personal communication); the oligochaete Tubifex hattai is an outgroup. Branch lengths do not reflect the genetic distance between taxa. Evolutionary changes of developmental characters are marked by short bars striking across the tree branches, and are assigned on the basis of parsimony. The role of the Q lineage as a P fate inducer evolved either as an apomorphy (1a) on the branch leading to the Helobdella leech species shown here, or is a plesiomorphic trait lost (1b) on the branch leading to Tubifex. The role of mesoderm as a redundant P fate inducer appears to have evolved as an apomorphy (2) on the branch leading to H. robusta (Austin) following its separation from H. robusta (Sacramento).





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