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Rmrs 2013 Culumber C001.Pdf RESEARCH Wide-Scale Population Sampling Identifies Three Phylogeographic Races of Basin Wildrye and Low-Level Genetic Admixture with Creeping Wildrye C. Mae Culumber, Steven R. Larson,* Thomas A. Jones, and Kevin B. Jensen C. Mae Culumber, Plants, Soils, and Climate Dep., Utah State Univ., Abstract Logan, UT 84322-4820; S.R. Larson, T.A. Jones, and K.B. Jensen, Basin wildrye [Leymus cinereus (Scribn. & Merr.) USDA-ARS, Forage and Range Research Lab., Utah State Univ., Á. Löve] and creeping wildrye [Leymus triti- Logan, UT 84322-6300. Received 29 June 2012. *Corresponding coides (Buckley) Pilg.] are outcrossing peren- author ([email protected]). nial grasses native to western North America. Abbreviations: AFLP, amplified fragment length polymorphism; These divergent species are generally adapted AMOVA, analysis of molecular variance; PCA, principal component to different habitats but can form fertile hybrids. analysis; PCR, polymerase chain reaction; QTL, quantitative trait Cultivars of both species are used in agricul- locus; UPGMA, unweighted pair group method with arithmetic mean. ture and conservation, but little is known about genetic diversity and gene flow among these species. Therefore, multilocus amplified frag- asin wildrye (Leymus cinereus) and creeping wildrye (Leymus ment length polymorphism (AFLP) genotypes Btriticoides) are closely related perennial grass species (Culumber and chloroplast DNA sequences were evaluated et al., 2011). Caespitose L. cinereus is considered the largest native from 536 L. cinereus and 43 L. triticoides plants grass in western North America, growing up to 3 m in height, and from 224 locations of western United States and is common throughout much of western United States and Canada Canada. Bayesian-cluster analysis detected (Barkworth, 2007). Leymus triticoides is a shorter, strongly rhizoma- three L. cinereus races corresponding to the tous species often associated with saline meadows, but its popula- Columbia, Rocky Mountain, and Great Basin regions. Possible admixture between species tions are less common and somewhat more restricted to California was detected in specific areas, but only 2.2% and the Great Basin region of Nevada, Utah, and southeast Ore- of the plants showed more than 10% introgres- gon (Barkworth, 2007). Both species are highly self-incompatible sion. The Columbia and Great Basin races were (Jensen et al., 1990) and known to form fertile hybrids (Dewey, predominantly octoploid whereas most of the 1970; Wu et al., 2003; Barkworth, 2007). Moreover, it has been Rocky Mountain accessions were tetraploid. suggested that the occasional presence of rhizomes in some speci- Approximately 36 and 7% of the AFLP variation mens of L. cinereus may be the result of introgression from these was apportioned among species and races, interspecific hybrids (Barkworth, 2007). In its simplest form, Ley- respectively, but no discrete marker differences mus Hochst. is allotetraploid (2n = 4x = 28), which is also true for were detected among these groups. Although both L. cinereus and L. triticoides. Octoploid (2n = 8x = 56) forms species can be distinguished using a relatively of L. cinereus and other Leymus species have also been reported small set of AFLP markers showing divergent (Barkworth, 2007), which may result from hybridization of diver- allele frequencies, at least 30 markers were needed to classify plants by race. Approximately gent species (Anamthawat-Jónsson and Bödvarsdóttir, 2001) or 8 and 11% of the chloroplast DNA sequence variation was apportioned among species and races, but these markers were not practically Published in Crop Sci. 53:996–1007 (2013). useful for species or race identification. doi: 10.2135/cropsci2012.06.0396 © Crop Science Society of America | 5585 Guilford Rd., Madison, WI 53711 USA All rights reserved. No part of this periodical may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or any information storage and retrieval system, without permission in writing from the publisher. Permission for printing and for reprinting the material contained herein has been obtained by the publisher. 996 WWW.CROPS.ORG CROP SCIENCE, VOL. 53, MAY–JUNE 2013 autoduplication. However, the geographic distribution of detected by molecular markers provides knowledge of pop- different ploidy levels within L. cinereus has not been docu- ulation structure resulting from historical effects of genetic mented and it is not known if the octoploids are genetically isolation, bottlenecks, and founder effects. Some markers different from tetraploids. may also be associated with allelic variation at quantita- Cultivars and germplasms of L. cinereus are used for tive trait loci (QTLs) controlling adaptive trait variation grazing, erosion control, and large-scale rangeland recla- (McKay and Latta, 2002). Experimental genetic mapping mation in western North America (Ogle, 2003). The culti- populations derived from interspecific hybrids of L. cinereus var Magnar (Alderson and Sharp, 1994) is believed to have and L. triticoides (Wu et al., 2003) have been used to iden- originated from a natural octoploid population in south- tify QTLs, genes, and DNA markers associated with adap- eastern British Columbia (Jones et al., 2009). The tetraploid tive traits such as plant height, caespitose and rhizomatous cultivar Trailhead was originally collected near Roundup, growth habits, and flowering traits (Larson et al., 2006), MT, and released in 1991 (Cash et al., 1998). Continental is fiber, protein, and mineral content (Larson and Mayland, a cultivar (Jones et al., 2009) derived from a chromosome- 2007), seed production, seed dispersal, and seed germina- doubled Trailhead population pollinated by the octoploid tion traits (Larson and Kellogg, 2009), and other function- Magnar, which shows increased seed mass and seedling ally important traits such as leaf glaucousness and gamete vigor compared to the parental cultivars. The L. cinereus compatibility (Larson et al., 2012). However, DNA markers germplasm Washoe (Marty 2003) was collected from a used to construct these maps (Wu et al., 2003) have not natural population growing on phytotoxic soils near the been tested on diverse natural populations and very little is now defunct Washoe smelter stack in the Anaconda Smelter known about the genetic diversity and gene flow within or Superfund Site in western Montana, which is contaminated between these two species. with As, Cd, Cu, Pb, and Zn (Marty, 2003). The L. triticoi- Phylogenetic analyses of chloroplast DNA sequences des cultivar Rio, released in 1991, was originally collected and multilocus nuclear AFLP genotypes demonstrated dis- in Kings Valley, CA, and is used for soil stabilization in tinctly different and polyphyletic relationships involving riparian areas, forage production, and reclamation of saline, North American and Eurasian Leymus species in compari- irrigated croplands and pasturelands (Young-Mathews sons with other Triticeae species, including wheat (Triticum and Winslow, 2010). Both L. cinereus and L. triticoides are aestivum L.) and barley (Hordeum vulgare L.) (Jones et al., well adapted to alkaline and saline soils that are common 1999; Redinbaugh et al., 2000; Liu et al., 2008; Zhou et al., throughout western North America and these plant mate- 2010; Culumber et al., 2011). In particular, it was shown that rials are used for a variety of agricultural and conservation the chloroplast genome of all Eurasian Leymus taxa are very purposes in this region (Ogle, 2003; Barkworth, 2007; similar to Psathyrostachys Nevski (Liu et al., 2008; Zhou et Young-Mathews and Winslow, 2010) including fire reha- al., 2010), which has been recognized as being the source of bilitation and other large-scale revegetation projects on fed- at least one of the diploid progenitors of allopolyploid Ley- eral lands, which total more than 182 million ha. mus (Dewey, 1970; Barkworth, 2007). Conversely, North Specific laws, regulations, and executive orders support American Leymus taxa, including L. cinereus and L. triticoides, the use of native plants on USDA Bureau of Land Manage- have a unique chloroplast genome that is more similar to ment and USDA Forest Service lands and there are impor- other Triticeae species including wheat and barley (Jones et tant concerns regarding the best seed sources for these al., 1999; Redinbaugh et al., 2000; Liu et al., 2008; Zhou native plants (Johnson et al., 2010). Similarities in environ- et al., 2010; Culumber et al., 2011). Presumably, the chlo- ment and climate between the site of plant-material origin roplast genome of North American Leymus comes from and target site or sites as well as the capacity to respond and an unknown progenitor species within this allopolyploid adapt to changing conditions are criterion used to deter- genus. Therefore, previous genetic studies of L. cinereus and mine plant material suitability (Conrad, 1983; McKay et L. triticoides and other Leymus taxa have revealed an interest- al., 2001; Johnson et al., 2004, 2010; Jones and Monaco, ing phylogeographic history of perennial grasses from North 2009). Moreover, strategies have been developed to increase America and Eurasia. However, previous studies were based genetic diversity of plant materials used for reclamation by on a very limited sampling of no more than six accessions sampling plants from multiple locations representing larger from any one species (Culumber et al., 2011). The objective genetic metapopulations,
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