“…Swensen (1996) addressed the question of homology of the symbioses in the four different actinorhizal groups by mapping anatomical and morphological features of root nodules onto the rbcL tree. These data included the number and shape of Frankia symbiotic vesicles (Baker and Schwintzer, 1990;Lalonde, 1979;Murray et al, 1985;Zhang et al, 1984), the location of Frankia vesicles within infected cells (Akkermans and van Dijk, 1981;Silvester and Harris, 1989), the location of infected cells within the nodule (Akkermans et al, 1983;Calvert et al, 1979;Newcomb and Pankhurst, 1982), evidence for gaseous diffusion barriers in the nodule and hemoglobin concentration (Tjepkema 1978;, and the mode of infection of the host plants (Berry et al, 1986;Berry and Sunell 1990; 1979; Kohls et al, 1994;Liu and Berry 1991;Miller and Baker, 1985;Racette and Torrey 1989a). There were clear differences in these characters in actinorhizal species belonging to each of the four groups, suggesting that groups B-E were the result of different symbiotic origins and that actinorhizal symbioses originated at least four times in the evolution of flowering plants.…”