December 2012 Number 1 - Utah Native Plant Society

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Utah Native Plant Society ated with the Mogollon Rim formation, but diploid populations appeared to be restricted to the western end of the rim. Hexaploid populations of P. amabilis were restricted to the easternmost portion of the range and appeared to be associated with magnesium rich igneous rock formations (data not shown). It is possible that such differences in distribution among cytotypes within species reflect ecological differentiation, which could result in local adaptation (Schonswetter et al. 2007; Buggs and Pannell 2007; Paun et al. 2007). Preliminary analysis of genetic variation in P. amabilis and P. woodhousei also supports differentiation among cytotypes (data not shown). We are continuing to investigate this unexpected variation in cytotype in both species using insights from genetics and ecology. CONCLUSIONS The results of this survey of cytotype variation among these western Phlox taxa demonstrate the value of such knowledge for the study of plant diversity, evolution, and conservation. The use of flow cytometry allowed us to gather data easily and rapidly on cytotype variation across expanded sample sizes that included multiple species from multiple locations throughout their ranges as well as population level sampling in species of conservation interest. We found that cytotype was variable not only among species but also among populations within species. This variation appears to be related to taxonomy, geography, and possibly ecology. Continued studies of cytotype variation in Phlox will provide valuable data for resolving taxonomic divisions as well as insight into the evolutionary diversification of this group. The detailed analysis of cytotype variation in P. amabilis and P. woodhousei, combined with results from ongoing population genetic work, suggest the presence of unique evolutionary lineages and ecological differentiation, both of which are important when planning conservation strategies. Taken together with recent reviews, these results emphasize the value of incorporating an understanding of cytotype variation into conservation biology studies. ACKNOWLEDGEMENTS We gratefully acknowledge support from the KSU Ecological Genomics Institute, the KSU Center for the Understanding of Origins and NSF DEB-0089656. We thank Thelma Green and Maureen Ty for technical assistance, Jerry Davison and Suzi Strakosh for assistance with the initial stages of flow cytometry, and Mark Mayfield and Ed Turcotte for unpublished chromosome count data. We thank the New York Botanical Garden (NY) for a loan of herbarium specimens. We thank Montezuma Castle National Monument (MOCA-2008- SCI-0004), Grand Canyon National Park (GRCA-2008- SCI-0015), the National Forest Service (various permits, Regions 1-6), the California BLM, and the Navajo Nation (Permits 414 and 051402-062) for permission to collect plants. This is publication 09-372-A of the Kansas Agricultural Experiment Station. LITERATURE CITED Bino, R.J., S. Lanteri, H.A. Verhoeven, and H.L. Kraak. 1993. Flow cytometric determination of nuclear replication stages in seed tissues. Annals of Botany 72: 181-187. Booy, G., R.J.J. Hendriks, M.J.M. Smulders, J.M. Van Groenendael, and B. Vosman. 2000. Genetic diversity and the survival of populations. Plant Biology 2: 379-395. Buggs, R.J.A. and J.R. Pannell. 2007. Ecological differentiation and diploid superiority across a moving ploidy contact zone. Evolution 61: 125-140. Cronquist A., A.H. Holmgren, N.H. Holmgren, J.L. Reveal, and P.K. Holmgren. 1984. Intermountain Flora, Vol. 4. New York Botanical Garden, Bronx, New York. Davison, J., A. Tyagi, and L. Comai. 2007. Largescale polymorphism of heterochromatic repeats in the DNA of Arabidopsis thaliana. BMC Plant Biology 7: 44. De Lange, P.J., P.B. Heenan, D.J. Keeling, B.G. Murray, R. Smissen, and W.R. Sykes. 2008. Biosystematics and conservation: a case study with two enigmatic and uncommon species of Crassula from New Zealand. Annals of Botany 101: 881-99. Dolezel, J. and J. Bartos. 2005. Plant DNA flow cytometry and estimation of nuclear genome size. Annals of Botany 95: 99-110. Ellis, J.R. and J.M. Burke. 2007. EST-SSRs as a resource for population genetic analyses. Heredity 99: 125-132. Ferguson, C.J. and R.K. Jansen. 2002. A chloroplast DNA phylogeny of eastern Phlox (Polemoniaceae): implications of congruence and incongruence with the ITS phylogeny. American Journal of Botany 89: 1324-1335. Ferguson, C.J., F. Krämer, and R.K. Jansen. 1999. Relationships of eastern North American Phlox (Polemoniaceae) based on ITS sequence data. Systematic Botany 24: 616-631. Ferguson, C.J., S.C. Strakosh, and R. Patterson. 2012. Phlox. Pp. 1068-1071 In: B. G. Baldwin, D.H. Goldman, D.J. Keil, R. Patterson, T.J. Rosatti, and D.H. Wilken, eds. The Jepson Manual, Vascular Plants of California, second edition. University of California Press, Berkeley. Grant, V. 1959. Natural history of the Phlox family: systematic botany. Martinus Nijhoff, The Hague, Netherlands. Halverson, K., S.B. Heard, J.D. Nason, and J.O. Stireman III. 2008. Origins, distribution, and local cooccurrence of polyploidy cytotypes in Solidago altis- 194
Calochortiana December 2012 Number 1 sima (Asteraceae). American Journal of Botany 95: 50- 58. Kron, P., J. Suda, and B.C. Husband. 2007. Applications of flow cytometry to evolutionary and population biology. Annual Review of Ecology, Evolution, and Systematics 38: 847-876. Levin, D.A. and B.A. Schaal. 1970. Reticulate evolution in Phlox as seen through protein electrophoresis. American Journal of Botany 57: 977-987. Levin, D.A. and D.M. Smith. 1966. Hybridization and evolution in the Phlox pilosa complex. The American Naturalist 100: 289-301. Löve, A. 1971. IOPB chromosome number reports XXXI. Taxon 20:157-160. Murray, B.G. and A.G. Young. 2001. Widespread chromosome variation in the endangered grassland forb Rutidosis leptorrhynchoides F. Muell. (Asteraceae: Gnaphalieae). Annals of Botany 87: 83-90. Paun, O., F.M. Fay, D.E. Soltis, and M.W. Chase. 2007. Genetic and epigenetic alterations after hybridization and genome doubling. Taxon 56: 649-656. Schonswetter, P., M. Lachmayer, C. Lettner, D. Prehsler, S. Rechnitzer, D.S. Reich, M. Sonnleitner, I. Wagner, K. Huelber, G.M. Schneeweiss, P. Travnicek, and J. Suda. 2007. Sympatric diploid and hexaploid cytotypes of Senecio carniolicus (Asteraceae) in the Eastern Alps are separated along an altitudinal gradient. Journal of Plant Research 120: 721-725. Severns, P.M. and A. Liston. 2008. Intraspecific chromosome number variation: a neglected threat to the conservation of rare plants. Conservation Biology 22: 1641-1647. Soltis, D.E., P.S. Soltis, D.W. Schemske, J.F. Hancock, J.N. Thompson, B.C. Husband, and W.S. Judd. 2007. Autopolyploidy in angiosperms: have we grossly underestimated the number of species? Taxon 56: 13- 30. Suda, J., P. Kron, B.C. Husband, P. Travnicek. 2007. Flow cytometry and ploidy: applications in plant systematics, ecology and evolutionary biology. In: J. Dolezel, J. Greilhuber, and J. Suda, eds. Flow Cytometry with Plant Cells: Analysis of Genes, Chromosomes and Genomes. Wiley-VCH, Weinheim. Pp. 103-130. Wherry, E.T. 1955. The genus Phlox. Morris Arboretum Monographs 3. Philadelphia, Pennsylvania. 195
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December 2012 Number 1 - Utah Native Plant Society

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