December 2012 Number 1 - Utah Native Plant Society

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Utah Native Plant Society 4 Family 7 4 Family 11 4 Family 14 4 Family 16 2 2 2 2 0 0 0 0 -2 -2 -2 -2 -4 -6 -4 -2 0 2 4 4 2 Family 17 -4 -6 -4 -2 0 2 4 4 2 Family 21 -4 -6 -4 -2 0 2 4 4 2 Family 24 -4 -6 -4 -2 0 2 4 4 2 Family 26 PRINCIPAL COMPONENT 2 0 -2 -4 -6 -4 -2 0 2 4 4 2 0 -2 4 2 0 -2 Family 27 -4 -4 -6 -4 -2 0 2 4 -6 -4 -2 0 2 4 Family 39 -4 -6 -4 -2 0 2 4 0 -2 -4 -6 -4 -2 0 2 4 4 2 0 -2 4 2 0 -2 Family 33 Family 41 -4 -6 -4 -2 0 2 4 0 -2 -4 -6 -4 -2 0 2 4 4 2 0 -2 Family 35 -4 -6 -4 -2 0 2 4 4 2 0 -2 Family 42 -4 -6 -4 -2 0 2 4 PCA 2 0 -2 -4 -6 -4 -2 0 2 4 4 2 0 -2 Family 37 -4 -6 -4 -2 0 2 4 One Individual Two Individuals Three Individuals Four Individuals Five Individuals PRINCIPAL COMPONENT 1 Figure 4. Scores on the first two axes from Principal Components Analysis of AFLP (amplified fragment length polymorphism) data for greenhouse-grown individuals belonging to 15 half-sib familes collected from the P. argillacea Tucker population in 2004. Point size reflects number of individuals with identical AFLP genotypes. argillacea indicated that individuals tended to group together by year much more than by population. These data suggest a persistent seed bank, which means a fraction of the seeds not only remain in the soil, but are viable for at least one year after production (Thompson and Grime 1979). A site characterization study of P. argillacea supported this hypothesis by suggesting that the seed bank reservoir contains an accumulation of seeds from many different years (Armstrong 1992). Preliminary results from a long term seed retrieval study with P. argillacea also support the existence of a longlived seed bank in this species. Few or no seeds have germinated in the field during the first two years, and most are still in a state of primary dormancy (Meyer unpublished data). This type of seed bank structure has also been reported in Phacelia secunda. Seeds from P. secunda were collected and allowed to germinate, and after three years a considerable fraction of the seeds remained viable but ungerminated (Cavieres 2001). PCA also suggests that collections from a single year and population of P. argillacea would depict a very narrow genetic diversity within the organism. A persistent seed bank can function as a genetic memory by accumulating seed genotypes from different years (Cabin et al. 1998). In the case of the rare annual Clarkia springvillensis, analysis of seed bank samples illustrated significantly higher within-seed bank genetic diversity when compared to the adult population (McCue and Holtsford 1998). The same could be true for P. argillacea, as evidenced by the pattern seen in the PCA (Figure 3). The seed bank must have a higher genetic diversity than the established plants in any one year, because of the wide range of diversity seen when comparing years. A similar situation was found in Phacelia dubia, which has small 132
Calochortiana December 2012 Number 1 population size and was thought to suffer from genetic drift and bottlenecking. However, analysis of the seed bank and adult plants from different years showed that no alleles were lost. The seed bank was found to store the full range of different genotypes (del Castillo 1994). Our study suggests that P. argillacea exhibits this type of age-structured seed bank and genetic pattern. In addition, the age and genotype of a seed may play a part in permitting it to germinate and establish in a particular kind of year. Allowing only certain genotypes to germinate each year would produce a pattern similar to the one found in Figure 3, with little or no overlap in genotypes between years. CONCLUSIONS In the current reintroduction efforts with P. argillacea, by selecting seeds collected in just one year we may be severely limiting the genetic base of this species. As the data from this study and similar studies suggest, in cases of a persistent seed bank, the parents of each year’s crop can differ from the seedling cohorts found in the years before and after (Figure 4). By using only greenhouse-grown seeds produced from the Tucker 2004 seed collection, we were inadvertently selecting for only a few specific genotypes. With individuals from just one year, the reintroduction program will almost surely suffer from inbreeding and genetic bottlenecks. To broaden the genetic base of this organism and allow for establishment of successful new populations of P. argillacea, the reintroduction program needs to include collections from several years and from both populations. ACKNOWLEDGMENTS This work was carried out with the aid of funding from the US Fish and Wildlife Service Preventing Extinction Program. We thank Heather Barnes (USFWS), Denise Van Keuren (formerly of the Uinta National Forest), Renee Van Buren and Kimball Harper (Utah Valley University), Katie Temus Merill and Ben Brulotte (Brigham Young University), Wendy Yates, Jennifer Lewinsohn, and Rita Dodge (Red Butte Gardens), Frank Smith (Western Ecological Services), and Bitsy Shultz and Susan Garvin Fitts (Utah Native Plant Society) for logistical assistance. Permission to collect tissue samples from specimens at the Brigham Young University Herbarium is also gratefully acknowledged. LITERATURE CITED Armstrong, V.R. 1992. Site characteristics and habitat requirements of the endangered Clay Phacelia (Phacelia argillacea Atwood, Hydrophyllaceae). M.S. thesis. Brigham Young University. Atwood, N.D. 1975. A revision of the Phacelia Crenulatae group (Hydrophyllaceae) for North America. Great Basin Naturalist 35: 1-190. Cabin, R.J., R.J. Mitchell, and D.L. Marshall. 1998. Do surface plant and soil seed bank populations differ genetically? A multipopulation study of the desert mustard Lesquerella fendleri (Brassicaceae). American Journal of Botany 85(9): 1098-1109. Cavieres, L.A. and M.T.K. Arroyo. 2001. Persistent soil seed banks in Phacelia secunda (Hydrophyllaceae): experimental detection of variation along an altitude gradient in the Andes of central Chile (33°S). Journal of Ecology 89: 31-39. del Castillo, R.F. 1994. Factors influencing the genetic structure of Phacelia dubia, a species with a seedbank and large fluctuations in population size. Heredity 72: 446-458. Garrison, L. 2007. Phylogenetic relationships in Phacelia (Boraginaceae) inferred from nrITS sequence data. M.S. thesis. San Francisco State University. Gilbert, C., J. Dempcy, C. Ganong, R. Patterson, and G.S. Spicer. 2005. Phylogenetic relationships within Phacelia subgenus Phacelia (Hydrophyllaceae) inferred from nuclear rDNA ITS sequence data. Systematic Botany 30: 627-634. Kang, M., Q. Ye, and H. Huang. 2005. Genetic consequence of restricted habitat and population decline in endangered Isoetes sinensis (Isoetaceae). Annals of Botany 96: 1265-1274. McCue, K.A. and T.P. Holtsford. 1998. Seed bank influences on genetic diversity in the rare annual Clarkia springvillensis (Onagraceae). American Journal of Botany 85: 30-36. Ronikier, M. 2002. The use of AFLP markers in conservation genetics – a case study on Pulsatilla vernalis in the Polish Lowlands. Cellular and Molecular Biology Letters 7: 677-684. Thompson, K and J.P. Grime. 1979. Seasonal variation in the seed banks of herbaceous species in ten contrasting habitats. Journal of Ecology 67: 893-921. U.S. Fish and Wildlife Service. 1978. Determination of five plants as endangered species. Federal Register 43: 44809-44812. U.S. Fish and Wildlife Service. 1989. Clay phacelia, Phacelia argillacea Atwood, recovery plan. USFWS Region 6, Denver Colorado. iii + 19 pp. Vos, P. et al. 1995. AFLP: a new technique for DNA fingerprinting. Nucleic Acids Research. 23: 4407-4414. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Higgins. 2003. A Utah Flora, third edition. Brigham Young University, Provo, UT. 133
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December 2012 Number 1 - Utah Native Plant Society

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