Phylogenetic, Morphological, and Chemotaxonomic Incongruence in the North American Endemic Genus Echinacea Lex Flagel Iowa State University

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Phylogenetic, Morphological, and Chemotaxonomic Incongruence in the North American Endemic Genus Echinacea Lex Flagel Iowa State University Ecology, Evolution and Organismal Biology Ecology, Evolution and Organismal Biology Publications 6-2008 Phylogenetic, Morphological, and Chemotaxonomic Incongruence in the North American Endemic Genus Echinacea Lex Flagel Iowa State University Ryan A. Rapp Iowa State University Corrinne E. Grover Iowa State University, [email protected] Mark P. Widrlechner United States Department of Agriculture, [email protected] Jennifer S. Hawkins UFonilvloerwsit ythi of sGe aondrgi additional works at: http://lib.dr.iastate.edu/eeob_ag_pubs Part of the Botany Commons, Ecology and Evolutionary Biology Commons, Molecular Genetics See next page for additional authors Commons, and the Plant Breeding and Genetics Commons The ompc lete bibliographic information for this item can be found at http://lib.dr.iastate.edu/ eeob_ag_pubs/60. For information on how to cite this item, please visit http://lib.dr.iastate.edu/ howtocite.html. This Article is brought to you for free and open access by the Ecology, Evolution and Organismal Biology at Digital Repository @ Iowa State University. It has been accepted for inclusion in Ecology, Evolution and Organismal Biology Publications by an authorized administrator of Digital Repository @ Iowa State University. For more information, please contact [email protected]. Authors Lex Flagel, Ryan A. Rapp, Corrinne E. Grover, Mark P. Widrlechner, Jennifer S. Hawkins, Jessie L. Grafenberg, Inés Álvarez, Gyu Young Chung, and Jonathan F. Wendel This article is available at Digital Repository @ Iowa State University: http://lib.dr.iastate.edu/eeob_ag_pubs/60 Botanical Society of America, Inc. Phylogenetic, morphological, and chemotaxonomic incongruence in the North American endemic genus Echinacea Author(s): Lex E. Flagel, Ryan A. Rapp, Corrinne E. Grover, Mark P. Widrlechner, Jennifer Hawkins, Jessie L. Grafenberg, Inés Álvarez, Gyu Young Chung and Jonathan F. Wendel Source: American Journal of Botany, Vol. 95, No. 6 (June 2008), pp. 756-765 Published by: Botanical Society of America, Inc. Stable URL: http://www.jstor.org/stable/27793066 Accessed: 07-10-2015 20:14 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/ info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Botanical Society of America, Inc. is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany. http://www.jstor.org This content downloaded from 129.186.176.217 on Wed, 07 Oct 2015 20:14:54 UTC All use subject to JSTOR Terms and Conditions American Journal of Botany 95(6): 756-765. 2008. PHYLOGENETIC, MORPHOLOGICAL, AND CHEMOTAXONOMIC INCONGRUENCE IN THE NORTH AMERICAN ENDEMIC GENUS EcHINACEA 1 LEX E. FLAGEL,2 RYAN A. RAPP,2 CORRINNE E. GROVER,2 MARK P. WIDRLECHNER,3 JENNIFER HAWKINS,4 JESSIE L. GRAFENBERG,2 INES ALVAREZ,5 GYU YOUNG CHUNG,6 AND JONATHAN F. WENDEL2,7 2Department of Ecology, Evolution, and Organismal Biology, Iowa State University, Ames, Iowa 50011 USA; 3USDA-ARS North Central Regional Plant Introduction Station, Ames, Iowa 5001 I USA; 4Department of Genetics, University of Georgia, Athens, Georgia 30602 USA; 5Departamento de Biodiversidad y Conservaci6n, Real Jardin Botanico, CSIC, Madrid 28014 Spain; and 6School of Bioresource Science, Andong National University, Andong Gyeongbuk, 760-749 South Korea The study of recently formed species is important because it can help us to better understand organismal divergence and the speciation process. However, these species often present difficult challenges in the field of molecular phylogenetics because the processes that drive molecular divergence can Jag behind phenotypic divergence. In the current study we show that species of the recently diverged North American endemic genus of purple coneflower, Echinacea, have low levels of molecular divergence. Data from three nuclear loci and two plastid loci provide neither resolved topologies nor congruent hypotheses about species-level relationships. This lack of phylogenetic resolution is likely due to the combined effects of incomplete lineage sorting, hybridiza­ tion, and backcrossing following secondary contact. The poor resolution provided by molecular markers contrasts previous studies that found well-resolved and taxonomically supported relationships from metabolic and morphological data. These results suggest that phenotypic canalization, resulting in identifiable morphological species, has occurred rapidly within Echinacea. Conversely, molecular signals have been distorted by gene flow and incomplete lineage sorting. Here we explore the impact of natural history on the genetic organization and phylogenetic relationships of Echinacea. Key words: Asteraceae; chloroplast DNA; Echinacea; incomplete lineage sorting; phylogenetics; single-copy nuclear DNA. Species of the genus Echinacea are geographically circum­ tantly, the disruption of natural processes of intraspecific and scribed within a region of North America that has undergone interspecific gene flow and the attendant increase in population repeated rounds of glaciation (Clayton and Moran, 1982), with fragmentation and genetic bottlenecks. the last such round, the Wisconsinan, ending roughly 10000 yr Taxonomically, Echinacea is delimited into nine species before the present. Presently, the genus ranges from southern (Table 1), including two, E. angustifolia DC and E. paradoxa Alberta, Canada to near the coast of the Gulf of Mexico in (Norton) Britton, that each are further divided into two varietals Texas and Louisiana and from the oak savannas of Ohio, the (McGregor, 1968; Flora of North America Editorial Commit­ glades of Tennessee, and open habitats in the Carolinas west to tee, 1993+; McKeown, 1999). These species are all diploid with the foothills of the Rocky Mountains (Urbatsch et al., 2006). the exception of E. pallida, which is putatively a polyploid Much of this range was under ice during the last glacial epoch, (Mechanda et al., 2004). This taxonomic treatment was devised signifying that Echinacea survived in southerly refugia. De­ by McGregor (1968), who spent 15 years studying the genus spite the expansive aggregate range of the genus, much of this while making controlled, common-garden crosses, noting that range has been converted into agricultural production, resulting many hybrids have high levels of stability, fecundity, and via­ in an extremely fragmented modern population structure. This bility in parental backcrosses. In a recent morphological study, distributional history has many potential implications for the four species with eight subspecies were proposed (Binns et al., genetic architecture of a perennial plant species, most impor- 2002, 2004), but McGregor's classification continues to be widely used by botanists and herbalists (see discussion in Blumenthal and Urbatsch [2006]) and serves as the basis for the 1 2008; revision accepted 7 April 2008. Manuscript received II February recent Flora of North America treatment (Urbatsch et al., 2006). paper of the Iowa Agriculture and Home Economics Experiment This journal regarding the ease of formation and fer­ Station, Ames, Iowa, Project No. 1018, was supported by Hatch Act and State McGregor's results ofiowa funds and was made possible by grant number P01ES012020 from the tility of interspecific hybrids suggest that Echinacea may either National Institute of Environmental Health Sciences (NIEHS) and the Office be a young genus in which rapid speciation has occurred of Dietary Supplements (ODS), NIH. lts contents are solely the responsibility (McKeown, 2004) or one in which, for reasons other than rela­ of the authors and do not necessarily represent the official views of the NIEHS, tive youth, genetic barriers have incompletely formed. In either NIH. Mention of commercial brand names does not constitute an endorsement case, gene flow between species has been historically common; of any product by the U. S. Department of Agriculture or cooperating agencies. McGregor noted hybrid swarms in natural sympatric settings, 1.-A. McCoy and other members of the Center for Research on Botanical and in more recent molecular work, Mechanda et al. (2004) in providing Dietary Supplements at Iowa State University were helpful found evidence of natural hybrid individuals. L. Urbatsch kindly provided feedback and comments throughout the research. and maintenance of Echinacea genetic diver­ DNA samples for sequence analysis. Assessment human health benefits 7 Author for correspondence (e-mail: [email protected]) sity is of interest due to the purported from several Echinacea species (Speroni et al., 2002), as well doi: 10.3732/ajb.0800049 as the cultivation and breeding of the plant as an ornamental 756 This content downloaded from 129.186.176.217 on Wed, 07 Oct 2015 20:14:54 UTC All use subject to JSTOR Terms and Conditions 757 June 2008] fLAGEL ET AL.-PHYLOGENETIC ANALYSIS OF THE GENUS ECHINACEA P. TABLE l. Echinacea taxa characterized, with U. S. state of origin and In contrast, a recent study (L. Wu, Iowa State University; USDA Germplasm Resources Infom1ation Network Plant Introduction Dixon, B. Nikolau, G. Kraus, M. Widrlechner, and E. Wurtele, (PI) accession."·b unpublished manuscript) of 40
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