Recent natural hybridization between two allopolyploid wheatgrasses (Elytrigia, Poaceae): ecological and evolutionary implications

BACKGROUND AND AIMS: Natural hybridization was investigated between two predominantly allohexaploid wheatgrasses, weedy Elytrigia repens and steppic E. intermedia, with respect to habitats characterized by different degrees of anthropogenic disturbance. METHODS: Using flow cytometry (relative DNA content), 269 plants from three localities were analysed. Hybrids were further analysed using nuclear ribosomal (ITS1-5.8S-ITS2 region) and chloroplast (trnT-F region) DNA markers in addition to absolute DNA content and chromosome numbers. KEY RESULTS: Weedy E. repens was rare in a steppic locality whereas E. intermedia was almost absent at two sites of agricultural land-use. Nevertheless, hybrids were common there whereas none were found at the steppic locality, underlining the importance of different ecological conditions for hybrid formation or establishment. At one highly disturbed site, > 16 % of randomly collected plants were hybrids. Hexaploid hybrids showed intermediate genome size compared with the parents and additive patterns of parental ITS copies. Some evidence of backcrosses was found. The direction of hybridization was highly asymmetric as cpDNA identified E. intermedia as the maternal parent in 61 out of 63 cases. Out of nine nonaploid cytotypes (2n = 9x = 63) which likely originated by fusion of unreduced and reduced gametes of hexaploids, eight were hybrids whereas one was a nonaploid cytotype of E. repens. The progeny of one nonaploid hybrid demonstrated gene flow between hexaploid and nonaploid cytotypes. CONCLUSIONS: The results show that E. repens and E. intermedia frequently cross at places where they co-occur. Hybrid frequency is likely influenced by habitat type; sites disturbed by human influence sustain hybrid formation and/or establishment. Hexaploid and nonaploid hybrid fertility is not negligible, backcrossing is possible, and the progeny is variable. The frequent production of new at least partially fertile cyto- and genotypes provides ample raw material for evolution and adaptation.     

Population studies in Goodyera (Orchidaceae) with emphasis on the hybrid origin of G. Tesselata

Morphological, cytological, and paper Chromatographic studies of populations from northern Michigan and examination of herbarium specimens from throughout North America were used to clarify the relationships ofGoodyera oblongifolia, G. repens var.ophioides, andG. tesselata. A canonical analysis of morphological data from mixed populations of these three species depictsG. tesselata as intermediate betweenG. oblongifolia andG. repens var.ophioides. The latter two species are diploid (2n = 30) andG. tesselata is tetraploid (2n = 60). Triploids (2n = ca. 45) were found in two mixed-species populations in northern Michigan.Goodyera tesselata produces three phenolic compounds present inG. oblongifolia and five different compounds present inG. repens var.ophioides. The range ofG. tesselata is confined to glaciated territory (except for two stations) in northeastern North America where the postglacially produced ranges ofG. oblongifolia andG. repens var.ophioides overlap. However,G. tesselata is quite abundant in areas outside the region of sympatry of the other two species. Based on this evidence, it is postulated thatG. tesselata is an allotetraploid species which resulted from hybridization betweenG. oblongifolia andG. repens var.ophioides during early post-Pleistocene. The slightly earlier blooming season ofG. tesselata may have been selected for to provide a measure of reproductive isolation between the tetraploid and its parents and to adapt the new species to the rather short growing season of northeastern North America.     

Genome size in Dahlia Cav. (Asteraceae–Coreopsideae)

The genus Dahlia (Asteraceae–Coreopsideae) is monophyletic according to a recent DNA phylogeny (ETS and ITS of rDNA). Traditionally, the genus has been divided into sections, but these have been shown not to be monophyletic. We have studied variation in genome size (DNA C-values) in a sample of species to investigate the possible effects of secondary metabolites on flow cytometry and Feulgen densitometry, and to see whether genome size variation has any systematic or phylogenetic significance. Using a range of cultivars, secondary compounds from corollas were shown to have only minor effects on the Feulgen method; the floral pigments were found to be relatively inert and seemed to have been extracted on fixation with acetic methanol. Freshly expanded corollas showed apparent apoptotic DNA decay in epidermal cells, so need to be used with caution. Flow cytometric measurements with propidium iodide in some cultivars resulted in a very similar average genome size (2C = 8.62 pg) as compared with Feulgen densitometry (2C = 8.84 pg). Leaf cytosol of D. variabilis has a demonstrable inhibitory effect on propidium iodide fluorescence, which may explain some of the intraspecific variation of C-values observed. DNA 2C-values ranged from 3.30 pg in D. dissecta (2n = 34) to 9.62 pg in a D. variabilis cultivar (2n = 64). The D. variabilis cultivars had broadly similar C-values showing a 1.16-fold range between cultivars. Some of this variation probably results from technical variables and the extent of genuine variation is uncertain. The highest 2Cx-value occurred in one D. coccinea accession (2.47 pg, 2n = 32; x = 8). D. coccinea with 2n = 64 showed slightly reduced Cx-values compared to D. coccinea with 2n = 32. Artificially produced interspecific hybrids had C-values that corresponded closely with expectations from the measured values obtained from their parents.     

Flow cytometric analysis reveals different nuclear DNA contents in cultivated Fonio (Digitaria spp.) and some wild relatives from West-Africa

Nuclear DNA amounts of 118 cultivated fonio accessions representing 94 landraces collected from the major growing areas of West-Africa (Benin, Burkina Faso, Guinea, Mali and Togo) and eight accessions of four wild relatives were investigated by Laser flow cytometry. In cultivated species, average 2C-values ranged from 1.848 ± 0.031 pg for Digitaria iburua to 1.956 ± 0.004 pg for D. exilis. In D. exilis landraces the chromosome number was determined at 2n = 36. The closely related wild species D. longiflora and D. ternata showed similar 2C DNA contents of 1.869 ± 0.035 pg and 1.775 ± 0.070 pg, respectively. Distinctly larger genomes were identified for more distant species D. lecardii and D. ciliaris with 2.660 ± 0.070 pg and 2.576 ± 0.030 pg per 2C nucleus, respectively. Intra-specific variations were found to be slight and insignificant, suggesting genome size stability mainly within the cultivated gene pool. These results support the distance of cultivated fonio species D. exilis and D. iburua from D. lecardii and D. ciliaris as well as their close relationships with D. longiflora and D. ternata. Relevance of the results for ploidy level considerations in fonio millets is discussed.     

Nuclear DNA C-values Complete Familial Representation in Gymnosperms

The gymnosperms are a monophyletic yet diverse group of woodytrees with approx. 730 extant species in 17 families. A recentsurvey showed that DNA C-values were available for approx. 16%of species, but for only 12 of the 17 families. This paper completesfamilial representation reporting first C-values for the fiveremaining families: Boweniaceae, Stangeriaceae, Welwitschiaceae,Cephalotaxaceae and Sciadopityaceae. C-values for nine Ephedraand two Gnetum species are also reported. C-values are now availablefor 152 (21%) species. Analysis confirms that gymnosperms arecharacterized by larger C-values than angiosperms (modal 1Cof gymnosperms = 15.8 pg compared with 0.6 pg in angiosperms)although the range (1C = 2.25–32.20 pg) is smaller thanthat in angiosperms (1C = 0.05–127.4 pg). Given completefamilial coverage for C-values and increasing consensus in gymnospermphylogeny, the phylogenetic component of C-value variation wasalso investigated by comparing the two datasets. This analysisrevealed that ancestral gymnosperms (represented by cycads and/orGinkgo; mean genome size = 14.71 pg) probably had larger genomes thanancestral angiosperms. Copyright 2001 Annals of Botany Company Gymnosperm DNA amounts, C-values, phylogeny, ancestral genome size, Cycadales, Ginkgo, Gnetales, conifers, Pinaceae     

Chromosome observations on some eastern North American Eleocharis (Cyperaceae)

Chromosome numbers are reported forEleocharis obtusa var.ellipsoidalis (2n = 10),E. obtusa var.peasei (2n =10),E. engelmannii (2n =10),E. olivacea f.olivacea (2n = 20),E. olivacea f.reductiseta (2n = 20),E. flavescens (2n = 30),E. tenuis var.verrucosa (2n = 20),E. tenuis var.tenuis (2n = 24),E. tenuis var.pseudoptera (2n = 39), a possible hybrid betweenE. tenuis var.pseudoptera and var.tenuis (2n = ca. 34 & 68),E. intermedia (2n = 22),E. tuberculosa (2n = 30),E. smallii (2n = 36), andE. ambigens (2n = 46). Noticeable chromosome size differences inE. tenuis, E. smallii, andE. ambigens are consistent with size differences previously reported by other authors. Variable meiotic pairing configurations were observed inE. flavescens, E. intermedia, andE. tenuis. Chromosome data help verify taxonomic relationships of some taxa and indicate the need for further investigation of others.     

Regeneration capacity from buds on roots and rhizomes in five herbaceous perennials as affected by time of fragmentation

Variation in seasonal sprouting pattern from roots and rhizomes of perennial herbaceous plants influence the success of plant proliferation ability, invasiveness and escape from weed control measures. The latter often rely on methods, which repeatedly fragment the underground system, thereby trigger adventitious and axillary buds to sprout, and consequently reduce the amount of stored energy. If carried out at times when no re-growth occurs, treatments will have little effect on weed populations, but cost much in terms of labour and energy. The purpose of this experiment was to determine the seasonal variation in bud sprouting capacity after fragmentation. Five troublesome perennial weed species, collected in northern and southern Sweden, were grown outdoors in Uppsala, Sweden (N 59°49′, E 17°39′), from May 2009 to January 2010. Cut root and rhizome fragments, taken at two weeks intervals from July to January, were used to evaluate bud sprouting capacity, which was statistically analyzed using generalized additive models. In Elytrigia repens from southern Sweden and Sonchus arvensis sprouting capacity was significantly impaired during a period from September to November. In Equisetum arvense and Tussilago farfara sprouting was low between July and November where after it increased. In contrast, Cirsium arvense and E. repens from northern Sweden sprouted readily throughout the period. Except for E. repens, a model by populations was significantly better than one based on latitudinal origin. The result suggests a species-specific timing of treatments in weed management, avoiding the non-effective autumn period for E. arvense, S. arvensis and T. farfara, and in some cases in E. repens.     

Consequence of ramet defoliation on plant clonal propagation and biomass allocation: Example of five rhizomatous grassland species

We investigated the effects of ramet defoliation frequency on clonal propagation and the patterns of biomass production and allocation on five rhizomatous species (Carex divisa Hude., Eleocharis palustris L., Juncus articulatus L., Juncus gerardii Lois. and Elytrigia repens L.). Plants were grown during an 18-week experiment in greenhouse conditions. The above ground parts of ramets were clipped following three treatments: frequent (every 2 weeks), moderate (every 4 weeks) and unclipped (control). The growth of C. divisa, J. articulatus and E. repens was strongly affected by defoliation whereas E. palustris and J. gerardii maintained a similar performance when defoliated. The latter were able to compensate for the biomass loss even after six consecutive clippings. Defoliation frequency had a significant effect on total biomass production for C. divisa, J. articulatus and E. repens while J. gerardii and E. palustris maintained total biomass production. Most of the studied species showed a decrease in clonal traits when defoliated. Clipped plants displayed fewer and shorter rhizomes. Defoliation had a strong influence in biomass production with a decrease in rhizome mass in all clipped species. A greater allocation to aerial parts and a lower to rhizomes were also detected. Moderate defoliation entailed intermediate response in 1/3 of detected significant effects of defoliation on plant traits. Finally, in the experimental conditions, E. palustris and J. gerardii were the most tolerant species to defoliation, while J. articulatus was intermediate and C. divisa and E. repens had the lowest tolerance.     

Nuclear DNA and heterochromatin contents in theScilla hohenackeri group,S. persica,andPuschkinia scilloides (Liliaceae)

DNA contents (presented as 1C-values) have been determined cytophotometrically in 7 species of theScilla hohenackeri group (10.18 to 22.71 pg), and in the systematically more isolated taxaS. persica (21.02 pg) andPuschkinia scilloides (6.80 pg). The heterochromatin amount is not correlated with the nuclear DNA content. Euchromatin, therefore, is not a particularly conservative part of the genome. However, high C-values and large but few terminal heterochromatin bands, and lower C-values and numerous but smaller heterochromatin bands are found to be linked in theS. hohenackeri group. Obviously, numerous chromosomal changes have accompanied speciation in this group. DNA contents, and C-banded karyotypes are consistent with systematic affinities based on morphological similarities.Evolution ofScilla and Related Genera, III.     

Polymorphic Sites in ITS1-5.8S rDNA-ITS2 Region in Hybridogenic Genus × <Emphasis Type="Italic">Elyhordeum</Emphasis> and Putative Interspecific Hybrids <Emphasis Type="Italic">Elymus</Emphasis> (Poaceae: Triticeae)

The genus Elymus L. is a complicated aggregate of ecological and geographical races, species, subspecies, varieties, and hybrids. We suggest that comparative analysis of intragenomic polymorphism of internal transcribed spacers ITS1 and ITS2 of 35S rRNA genes in the supposed hybrids and their possible “parents” can be one of the approaches to verification of hybrid origin of the samples collected in nature to confirm or reject the hypotheses about their possible “parents.” Polymorphic sites (PS) in ITS of 23 Elymus species, as well as in two supposed interspecific Elymus hybrids and in a supposed intergeneric hybrid between Elymus × Hordeum determined as × Elyhordeum sp., were analyzed in the work. We collected all hybrids in the Altai. There were 2 and 5 PS in two samples of E. dahuricus and 1 and 4 PS in two studied samples of E. schrenkianus in the ITS1-5.8S rDNA-ITS2 region. From 0 to 4 (modes 0 and 3) PS were detected in 32 samples relating to 21 tetraploid Elymus species. More PS (14) were found in the × Elyhordeum sp. sample. A large number of single nucleotide substitutions were found in 5.8S rRNA in × Elyhordeum. It was shown that about half of them do not change the secondary structure of the 5.8S rRNA molecule, so these molecules probably retain the ability to work as a component of large subunit of a ribosome. On the other hand, the absence or weakening of 5.8S rDNA homogenization in × Elyhordeum indirectly suggests that a significant part of 5.8S rDNA is not transcribed. Paradoxically, ITS sequences of × Elyhordeum sp. are less polymorphic than 5.8S rDNA. There are no ITS sequences derived from Hordeum among × Elyhordeum ITS sequenced by Sanger method. No traces of the H subgenome and a subgenome originating from Agropyron (P-subgenome) are seen in the Alt 10–278 plant genome (a chimera, combining the morphological traits of Elymus, Elytrigia, and Agropyron). In this plant, as well as in the supposed intersectional hybrid Alt 11–60 distinguished by a mosaic of the traits typical for the E. caninus × E. mutabilis species, only 4 and 5 PS, respectively, are detected when sequencing by Sanger method. The comparison of ITS sequences of the supposed Elymus Alt 10–278 hybrid and its probable “parents” demonstrates that one of the species of the Elymus macrourus kinship circle, as well as the Elytrigia geniculata, could be one of its ancestors. The comparison of the ITS sequence of the supposed parental species with ITS of Alt 11–60 samples and five PS of the supposed Alt 11–60 hybrid does not contradict the hypothesis that this is an intersectional hybrid of the first generation that emerged with the involvement of E. caninus and E. mutabilis common in the Altai.     

A first assessment of genome size diversity in Monogonont rotifers

This study provides the first analysis of genome size diversity in Monogonont rotifers. Measurements were made using flow cytometry, with Drosophila melanogaster and chicken erythrocytes as internal standards. Nuclear DNA content (??2C????assuming diploid genomes) in eight different species of four different genera ranged almost fourfold, from 0.12 to 0.46 pg. A comparison with previously published values for Bdelloid rotifers suggested that the genomes of Monogononts are significantly smaller than those of Bdelloids. When compared to other Metazoans, Monogonont rotifers seem to have relatively small genomes. For instance, the C-values of the two species with the smallest genomes, Brachionus dimidiatus and Synchaeta pectinata, were only 0.06 and 0.085 pg, respectively. Various explanations for genome size diversity within Monogononta are discussed.     

Compared chemical properties of dermonecrotic and lethal toxins from spiders of the genusLoxosceles (Araneae)

Loxosceles spider venom usually causes a typical dermonecrotic lesion in bitten patients, but it may also cause systemic effects that may be lethal. Gel filtration on Sephadex G-100 ofLoxosceles gaucho, L. laeta, orL. intermedia spider venoms resulted in three fractions (A, containing higher molecular mass components, B containing intermediate molecular mass components, and C with lower molecular mass components). The dermonecrotic and lethal activities were detected exclusively in fraction A of all three species. Analysis by SDS-PAGE showed that the major protein contained in fraction A has molecular weight approximately 35 kDa inL. gaucho andL. intermedia, but 32 kDa inL. laeta venom. These toxins were isolated from venoms ofL. gaucho, L. laeta, andL. intermedia by SDS-PAGE followed by blotting to PVDF membrane and sequencing. A database search showed a high level of identity between each toxin and a fragment of theL. reclusa (North American spider) toxin. A multiple sequence alignment of theLoxosceles toxins showed many common identical residues in their N-terminal sequences. Identities ranged from 50.0% (L. gaucho andL. reclusa) to 61.1% (L. intermedia andL. reclusa). The purified toxins were also submitted to capillary electrophoresis peptide mapping afterin situ partial hydrolysis of the blotted samples. The results obtained suggest thatL. intermedia protein is more similar toL. laeta toxin thanL. gaucho toxin and revealed a smaller homology betweenL. intermedia andL gaucho. Altogether these findings suggest that the toxins responsible for most important activities of venoms ofLoxosceles species have a molecular mass of 32–35 kDa and are probably homologous proteins.     

The Resistance of <Emphasis Type="Italic">Phleum pratense</Emphasis> and <Emphasis Type="Italic">Elytrigia repens</Emphasis> to High Concentrations of Zinc

The influence of increased zinc concentrations on seed germination, growth activity, photosynthetic apparatus, and water metabolism in two perennial grasses (Phleum pratense L. and Elytrigia repens (L.) Nevski) was studied in laboratory and vegetation experiments to assess plant metal tolerance. In laboratory conditions it was established that seeds of both species may germinate in a wide range of zinc concentrations. In vegetation experiments, the possibility of successful growth and accumulation of biomass of both grasses in the presence of high zinc concentration in the root medium was revealed. At the same time, high water contents in root and shoot tissues were maintained, as well as the necessary intensity of photosynthesis (due to maintenance of the efficiency of photosystem II and the amount of carotenoids). It was noted that the established high resistance of both species of grasses to zinc, as well as their ability to accumulate significant amounts of metal ions in the roots, indicates that P. pratense and E. repens may be used for phytoremediation of soils contaminated with zinc.     

Nuclear DNA content in two Sphaeriidae species from Southern Chile (Bivalvia: Veneroida)

The family Sphaeriidae is a group of cosmopolitan freshwater clams that inhabit temporary and permanent fluvial and lacustrine environments around the world. In Chile, Sphaeriidae is represented by 11 species belonging to the genera Musculium, Pisidium and Sphaerium. In this study, the nuclear DNA content (or 2C-value) of Musculium argentinum (d’Orbigny, 1835) and Pisidium llanquihuense Ituarte, 1999 was estimated for the first time. Musculium argentinum showed a 2C-value of 3.7?±?0.38?pg, whereas P. llanquihuense showed a 2C-value of 3.57?±?0.36?pg. These 2C-values are within the range previously reported for Sphaeriidae (2.35–5.7?pg), which includes diploids and polyploids, amounting to 13 of the species examined around the world. Of the cytological data available for the family, the nuclear DNA content can have a unifying role in understanding the magnitude of the chromosome polymorphisms within this polyploid complex.     

Taxonomy of the bouteloua repens complex

An investigation was made of grasses variously referred toBouteloua americana,B. filiformis, B. radicosa, B. repens, B. bromoides, B. heterostega, B. pubescens, andB. williamsii. Four of these species,B. americana, B. williamsii, B. repens, andB. radicosa, are maintained as separate taxa; the other four are reduced to synonymy underB. repens. Described as new isB. longiseta, with a tropical distribution from Oaxaca, Mexico, south to Honduras. Chromosome counts of 2n = 20 forB. williamsii and 2n = 40 forB. longiseta are the first for these taxa. Counts of 2n = 20, 2n = 40, and 2n = 60, and a few aneuploid numbers, are reported forB. repens     

Flow cytometry and cytogenetic tools in eucalypts: genome size variation × karyotype stability

The eucalypts comprise a group of woody plants used in commercial forest plantations owing to their high growth rates, adaptability to various ecological conditions and multiple applications. Despite the enormous amount of molecular data available for eucalypts, a basic understanding of the nature of its genome still requires information regarding the DNA amount in the genus. In this work, we estimated the genome size and base composition of 25 eucalypt species. With a comparative karyotype approach, we aimed to identify possible chromosomal alterations correlated with the genome size variation. Classical cytogenetic and genomic in situ hybridization experiments were conducted for this purpose. The studied species showed genome size ranging from 2C = 0.91 (Corymbia intermedia) to 2C = 1.37 pg (Eucalyptus paniculata) and AT/CG ratios varying from AT = 61.3 (Eucalyptus urophylla) to AT = 62.85% (C. intermedia). Comparative karyotype analysis revealed no remarkable differences in chromosome number (2n = 22) or morphology among eucalypt species despite considerable differences in nuclear DNA content. The genome in situ hybridization method did not distinguish non-homologous chromosomal regions of Eucalyptus baileyana and Corymbia citriodora, despite the difference of 0.45 pg between their genome sizes. The results found in the present work corroborate the consideration of small and dispersed DNA changes as the main cause of genome size variation in eucalypts.     

Effect of rotational and continuous grazing on vegetation of an upland grassland in the Jizerské hory Mts., Czech Republic

The effect of different grazing regimes on pasture vegetation was studied during the intensive grazing of heifers in the Jizerské hory mountains during 1993–1997. The vegetation was monitored in 3 pairs of permanent 1×1 m plots using a continuous grid of nine 0.33 × 0.33 m subplots. We applied continuous stocking and rotational grazing. Vegetation varied as a result of time and differences between treatments. Several prostrate dicotyledonous species (Trifolium repens, Taraxacum sp.,Bellis perennis andLeontodon autumnalis) increased under continuous stocking. This treatment also promoted the growth of the perennial grassLolium perenne, which was able to cope with frequent defoliation. Tall grasses sensitive to frequent defoliation (Poa trivialis, Holcus mollis, Alopecurus pratensis, Dactylis glomerata andElytrigia repens) were more abundant in rotationally grazed paddocks. Species diversity was not significantly influenced by the different grazing systems. The decrease in the potential sward height under continuous stocking revealed the replacing of tall dominants by lower species. Our results indicate that different grazing systems alter the composition and structure of grassland vegetation. Defining the intensity of grazing under continuous stocking or rotational grazing is complex due to the different stocking rates and the heights of sward during a grazing season. Information about pasture management should therefore involve not only grazing intensity but also the grazing system used.     

Influence of a type 2 diabetes associated prostaglandin E synthase 2 polymorphism on blood prostaglandin E2 levels

In this study we tested whether the type 2 diabetes mellitus associated prostaglandin E synthase 2 arginine to histidine polymorphism at position 298 (R298H) influences prostaglandin E2 levels in humans. Fasting prostaglandin E2 was determined in the blood of subjects carrying different genotypes of the R298H polymorphism. Subjects were matched by sex, age, and body mass index. No differences in prostaglandin E2 levels were found with respect to genotypes when considering the whole group. Male homozygous histidine carriers showed elevated prostaglandin E2 levels compared to heterozygous carriers and homozygous arginine carriers (188.2±42.4 vs. 80.4±26.5 pg/ml, p=0.021; and vs. 92.9±15.3 pg/ml, p=0.11). These differences were not evident in female subjects. In contrast, 6-keto-prostaglandin F1alpha levels as independent marker of arachidonic acid metabolism showed ambiguous results. Nevertheless, preliminary evidence of the prostaglandin E synthase 2 R298H polymorphism possibly influencing prostaglandin E2 blood levels in a gender-specific manner was obtained.     

A serological and disc electrophoretic study of North American Typha

Pollen and seed proteins of seven selected North American and Puerto RicanTypha populations were compared using two serological methods and disc electrophoresis. These methods were capable of discriminating among all taxa studied:Typha latijolia, T. angustifolia, T. ×glauca, andT. domingensis. The two hybrid populations were found to contain proteins not found in either parent.Typha domingensis was serologically the most distinct of the four taxa. The diagnostic morphological characteristics forTypha species were studied in all populations, and statistical comparisons are presented. Data from the morphological observations agreed with the information obtained from the chemosystematic research. All data indicate that the three taxa should be maintained as separate species. The hybrid nature of the putativeT. ×glauca is verified by both the biochemical and morphological data. Observed morphological and biochemical differences support taxonomic treatments in whichT. domingensis is designated as a separate species     

Hybridisation,genomic constitution and generic delimitation inElymus s. l. (Poaceae: Triticeae)

Intergeneric crosses were made between representatives of the genomically-defined generaElymus, Agropyron, Elytrigia, Pseudoroegneria, andThinopyrum. The genomic constitution ofElytrigia repens, the type species ofElytrigia, is shown to be SSH, a genomic combination otherwise found only inElymus. The S genome ofPseudoroegneria has almost always a dominant influence on the morphology of the taxa of which it is a component.Wang (1989) showed that the J genome inThinopyrum and the S genome have considerable homoeology, with a mean c-value of 0.35 in diploid SJ hybrids. A genetic coherence from S to SJ^e, J^e, J^eJ^b, and J^b can be expected, agreeing with the continuous morphologic variation pattern observed. Because of the absence of morphological discontinuities between the taxa,Pseudoroegneria (S),Elymus (SH, SY, sometimes with additional genomes),Elytrigia (SSH, SSHX), andThinopyrum (SJ, SJJ, J) are best treated as a single genus,Elymus, following the generic concept ofMelderis in Flora Europaea and Flora of Turkey. The basic genomic constituents ofElymus will then be the S and/or J genomes.Agropyron, with diploids, tetraploids, and hexaploids based on the P genome is morphologically distinct from other genera inTriticeae. In a few species ofElymus andPseudoroegneria, a P genome is an additional constituent. In these cases the P genome has a negligible morphological influence. Therefore, it seems reasonable to maintainAgropyron as a separate genus.