A space‐for‐time substitution reveals the long‐term decline in genotypic diversity of a widespread salt marsh plant, Spartina alterniflora, over a span of 1500 years

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Abstract
Summary: Clonal populations face a trade‐off between sexual recruitment and vegetative growth and, once established, may undergo continuous declines in genotypic diversity if their sexual recruits make poor competitors. The geological history of delta formation in the Lower Mississippi River Valley was used to age eight S. alterniflora marshes for use in a space‐for‐time substitution ranging over 1500 years, in order to determine the long‐term effects of clonal growth on genotypic diversity in natural populations. We also predicted that highly heterozygous clones are competitively superior, leading to an increase in the overall level of genetic diversity as a marsh ages and/or to an increasingly positive relationship between clone size and individual heterozygosity, and that the clumping of ramets within clones will occur over increasingly large distances as populations age, while the clumping of genetically related clones will become less pronounced as intraclonal competition begins to obscure the initial effects of localized seedling recruitment. Using molecular markers to differentiate clones, we documented a decline in clonal richness at the rate of approximately 1% 100 years−1 that was accompanied for the first 300–500 years by an increase in the distance over which clumping of ramets within genets occurred. Older populations, in the 500–1500‐year range, showed evidence of clone fragmentation. The spatial clustering of kin was observed for only two marshes, and exhibited no clear relationship with marsh age. Whereas the overall level of genetic diversity was consistent among marshes and showed no clear relationship with marsh age, the relationship between heterozygosity and individual clone size became increasingly pronounced within older marshes. Our results suggest that under natural conditions S. alterniflora marshes will rarely reach ages sufficient for the loss of all clonal diversity, or for the effects of inbreeding and drift to pose a significant threat to population viability.