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Allopolyploids of the Genus Elymus (Triticeae, Poaceae): a Phylogenetic Perspective Roberta J

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Allopolyploids of the Genus Elymus (Triticeae, Poaceae): a Phylogenetic Perspective Roberta J Aliso: A Journal of Systematic and Evolutionary Botany Volume 23 | Issue 1 Article 30 2007 Allopolyploids of the Genus Elymus (Triticeae, Poaceae): a Phylogenetic Perspective Roberta J. Mason-Gamer University of Illinois, Chicago Follow this and additional works at: http://scholarship.claremont.edu/aliso Part of the Botany Commons, and the Ecology and Evolutionary Biology Commons Recommended Citation Mason-Gamer, Roberta J. (2007) "Allopolyploids of the Genus Elymus (Triticeae, Poaceae): a Phylogenetic Perspective," Aliso: A Journal of Systematic and Evolutionary Botany: Vol. 23: Iss. 1, Article 30. Available at: http://scholarship.claremont.edu/aliso/vol23/iss1/30 Aliso 23, pp. 372–379 ᭧ 2007, Rancho Santa Ana Botanic Garden ALLOPOLYPLOIDS OF THE GENUS ELYMUS (TRITICEAE, POACEAE): A PHYLOGENETIC PERSPECTIVE ROBERTA J. MASON-GAMER University of Illinois at Chicago, Department of Biological Sciences, MC 066, 845 West Taylor Street, Chicago, Illinois 60607, USA ([email protected]) ABSTRACT The wheat tribe, Triticeae, includes many genomically distinct polyploid taxa. Elymus is an entirely allopolyploid genus, with all species containing the St genome of Pseudoroegneria. The St genome may be combined with one or more distinct genomes representing multiple, diverse diploid donors from throughout the tribe. This study includes a simultaneous phylogenetic analysis of new and pre- viously published data from several distinct Elymus groups, including North American and Eurasian StStHH tetraploids, in which the H genome is derived from Hordeum, Eurasian StStYY tetraploids, in which the Y genome is derived from an unknown donor, and a putative StStStStHH hexaploid. Elymus species were analyzed with a broad sample of diploid genera from within the tribe using a combination of molecular data from the chloroplast and the nuclear genomes. The data confirm the genomic constitution of the StStHH and StStYY tetraploids, but do not provide additional information on the identity of the Y-genome donor. The genomic diversity in the hexaploid is greater than expected, inconsistent with the hypothesis of an StStStStHH genome complement. Key words: Elymus, Poaceae, polyploidy, reticulation, Triticeae, wheat tribe. INTRODUCTION diploid and autotetraploid species, which are classified as Pseudoroegneria (Dewey 1984). In Elymus, the St genome The wheat tribe, Triticeae (Pooideae, Poaceae), is widely can be combined with genomes from one or more Triticeae known for its economic importance. This monophyletic tribe genera, including Hordeum L. (genome designation H), Ag- includes three major grain crops—wheat, barley, and rye— ropyron Gaertn. (P), Australopyrum (Tzvelev) A´ . Lo¨ve (W), along with several important forage grasses and numerous and an unknown donor (Y), in various allopolyploid com- weedy, invasive species. From an evolutionary standpoint, binations including StStHH, StStYY, StStHHHH, StSt- the tribe’s complex reticulate history has been of interest for StStHH, StStYY, StStStStYY, StStYYYY, StStHHYY, many years. At the diploid level, conflict among gene trees, StStYYWW, and StStYYPP (e.g., Dewey 1967, 1968, especially between those based on chloroplast and nuclear 1970b, 1974, 1984; Jensen 1990, 1993, 1996; Salomon and DNA data (Mason-Gamer and Kellogg 1996a), suggests a Lu 1992, 1994; Lu and von Bothmer 1993; Lu et al. 1995). history of gene exchange, lineage sorting, or a combination Other St-containing allopolyploids include Pascopyrum of both (Kellogg et al. 1996). Furthermore, polyploidy is smithii (Rydb.) Barkworth & D. R. Dewey, which combines common in the tribe; about 75% of the species with known the Pseudoroegneria and Hordeum genomes with the Ns ge- chromosome numbers are polyploid (Lo¨ve 1984). nome of Psathyrostachys Nevski in an StStHHNsNsNsNs Allopolyploidy represents a reticulate process by which octoploid configuration (Dewey 1975), and Thinopyrum A´ . distinct genomes are united in a single nucleus. While there Lo¨ve, some species of which are hypothesized to combine are problems with placing reticulate taxa within the bifur- cating trees that are obtained using many methods of anal- the St genome with the E and/or J genomes usually consid- ysis (e.g., Hull 1979; Cronquist 1987; McDade 1992, 1998), ered characteristic of Thinopyrum (e.g., Liu and Wang 1993; studies of individual gene trees often allow these problems Zhang et al. 1996; Chen et al. 1998). Thus, the St genome, to be circumvented. Thus, molecular phylogenetic data have probably more than any other in Triticeae, plays an impor- recently revealed the reticulate histories of several polyploid tant role in the complex reticulate allopolyploid patterns that species or groups, for example: Gossypium L. (e.g., Cronn characterize the tribe. et al. 1996, 2003; Seelanan et al. 1997; Small and Wendel This overview focuses on a subset of species from the 2000), Geum L. (Smedmark et al. 2003), Glycine Willd. genomically heterogeneous Elymus, and presents molecular (e.g., Doyle et al. 2002; Rauscher et al. 2002), Oxalis L. phylogenetic data that address their relationships to the dip- (Emshwiller and Doyle 1998, 2002), Oryza L. (Ge et al. loid members of the tribe. These species include (1) North 1999), and Paeonia L. (e.g., Sang and Zhang 1999). American tetraploid species with presumed StStHH genome Within Triticeae, Elymus L. is a genomically heteroge- configurations (e.g., Dewey 1982, 1983a, b, 1984; Mason- neous polyploid group of about 140 species (Lo¨ve 1984). Gamer 2001; Mason-Gamer et al. 2002; Helfgott and Ma- According to its genomic definition, Elymus comprises al- son-Gamer 2004), (2) Eurasian tetraploid species, with pre- lopolyploid species with at least one set of chromosomes sumed StStHH or StStYY genome configurations (e.g., Lu derived from Pseudoroegneria (Nevski) A´ . Lo¨ve (genome and von Bothmer 1990, 1993; Salomon and Lu 1992, 1994; designation St). The St genome can also be found in both Lu 1993; Lu et al. 1995), and (3) a Eurasian hexaploid spe- VOLUME 23 Elymus Phylogeny 373 cies (E. repens L.), with a presumed StStStStHH genomic sequence. Gaps in the sequence alignments were treated as configuration (Cauderon and Saigne 1961; Dewey 1970a, missing data. Branch support was estimated using 1000 ‘‘fast 1976; Ørgaard and Anamthawat-Jo´nsson 2001). These gen- bootstrap’’ replicates, as implemented in PAUP* vers. omically diverse allopolyploids are analyzed together with a 4.0b10 (Swofford 2002). Trees were rooted using Psathy- broad sample of diploid genera from throughout the tribe, rostachys; two earlier cpDNA analyses (Mason-Gamer and using three sources of chloroplast DNA (cpDNA) data, and Kellogg 1996a; Petersen and Seberg 1997) used Bromus L. sequences from the single-copy nuclear gene encoding gran- (tribe Bromeae) as an outgroup and both studies provided ule-bound starch synthase (GBSSI). strong evidence (100% bootstrap in both cases) that Psathy- rostachys is at the base of the Triticeae cpDNA tree. MATERIALS AND METHODS The GBSSI data set adds newly acquired data from five tetraploid Eurasian species to existing data from represen- Both of the molecular data sets include a combination of tatives of most of the monogenomic genera of the tribe (Ma- new and previously published data. Specimens correspond- son-Gamer et al. 1998; Mason-Gamer 2001), from six North ing to the new data are currently in the possession of the American StStHH tetraploids (E. elymoides, E. glaucus, E. author, and herbarium vouchers will be permanently depos- lanceolatus, E. trachycaulus, E. virginicus, and E. wawa- ited upon the completion of the ongoing study. Information waiensis; Mason-Gamer 2001) and from hexaploid E. repens about the previously published sequence data and the plants (Mason-Gamer 2004). New data were obtained from E. from which they were derived can be found in the corre- abolinii (China; PI531555), E. caninus L. (Uzbekistan; sponding publications, cited below. PI314205), E. ciliaris (China; PI531575), E. dentatus (Rus- The cpDNA data set includes restriction sites, sequences sia; PI628702), and E. mutabilis (Russia; PI628704). A 1.3 from trnT, trnL, and trnF genes and their intergenic spacers, kilobase portion of the GBSSI gene was amplified, cloned, and sequences of the rpoA gene. The existing restriction site and sequenced as in Mason-Gamer (2001) using primers F- data set (Mason-Gamer and Kellogg 1996b) includes most for, H-for, J-bac, L1-for, L2-bac, and M-bac (Mason-Gamer of the monogenomic genera and three species of Elymus (E. et al. 1998). Because multiple, distinct nuclear sequences are lanceolatus (Scribn. & J. G. Sm.) Gould, E. repens, and E. expected to coexist in allopolyploid individuals, multiple ciliaris (Trin.) Tzvelev); no new restriction site data were clones per individual (4–12 from each tetraploid, 20ϩ from added for the present analysis. The existing tRNA/spacer and E. repens) were partially sequenced and compared in an at- rpoA data sets include most of the monogenomic genera tempt to obtain, and then fully sequence, all variants within (Mason-Gamer et al. 2002), eight North American StStHH each individual. The ten new tetraploid sequences included tetraploids (E. canadensis L., E. elymoides (Raf.) Swezey, here have been assigned GenBank accession numbers E. glaucus Buckley, E. hystrix L., E. lanceolatus, E. trachy- DQ159322–DQ159331. caulus (Link) Gould ex Shinners, E. virginicus L., and E. The GBSSI data were analyzed using PAUP* vers. 4.0b10 wawawaiensis J. Carlson & Barkworth; Mason-Gamer et al. (Swofford 2002). Initial cladistic
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