CHROMOSOMAL ANALYSES AND THE ORIGIN OF ALLOPOLYPLOID POLYPODIUM VIRGINIANUM (POLYPODIACEAE)

Polypodium virginianum L. has been treated as a morphologically and chromosomally variable species. Recently, however, two species have been named from within the P. virginianum complex. Chromosome counts reported here confirmed that these segregate species, P. appalachianum Haufler and Windham and P. sibiricum Siplivinskij, are diploids with n = 37. It was further hypothesized that hybridization between these diploids initiated the allotetraploid P. virginianum. Naturally occurring, sterile triploid plants obtained from habitats in which the putative allotetraploid was sympatric with its presumed diploid progenitors have been verified isozymically as backcross hybrids. Meiotic chromosomal behavior showed that the triploids contained univalents and bivalents in nearly equal numbers, confirming that tetraploid P. virginianum contained one genome from each diploid. Observation of three to six trivalents in triploid preparations suggested that the diploids are still closely related genetically. This hypothesis was supported by isozymic studies which found a genetic identity of 0.610 between P. appalachianum and P. sibiricum, a value much higher than the average identity for congeneric fern species studied to date (0.33) and higher than any other species pair in the P. vulgare L. complex (average I = 0.307). These chromosomal and isozymic data help to resolve a biosystematic riddle that has interested pteridologists for over 40 years.