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    Patterns of Genetic Variation in Southern Appalachian Populations of Athyrium filix‐femina var. asplenioides (Dryopteridaceae)

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    Allozyme variation (17 loci coding 11 enzymes) was investigated in 14 populations of the fern Athyrium filix‐femina var. asplenioides arrayed at differing elevations and latitudes in the southern Appalachians. Allozyme fingerprints showed that asplenioides individuals comprise meandering, overlapping clones usually ≤3 m in extent, occasionally forming larger clones of up to 17 m. Levels of genetic variability in populations (means: , , ) were near the averages for both ferns and seed plants. General conformance to Hardy‐Weinberg expectations indicated a predominantly outcrossing mating system. Hierarchical F statistic analysis and occasional deficits and excesses of heterozygotes indicated population substructure. Similar allele frequencies across all populations resulted in low to moderate values (mean ; ) and high values of genetic similarity (mean ; mean ). Hierarchical analysis indicated that neither regional proximity ( ) nor elevation ( ) contributed substantially to divergence among populations ( ), a result corroborated by UPGMA analysis that clustered together populations from different regions and of different elevational class. Southern Appalachian asplenioides differed from more eastern asplenioides populations of the piedmont and coastal plain in having higher frequencies of Pgm‐2c and Tpi‐2B, alleles characteristic of the more northern variety angustum. Nonetheless, genetic distinctness of the two varieties was maintained. We hypothesize that higher frequencies of angustum alleles in the southern Appalachian asplenioides populations are the result of introgression from angustum that persisted at high elevations as both taxa migrated northward following the retreat of the Wisconsinan glacier
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