Competition among native and invasive Phragmites australis populations: An experimental test of the effects of invasion status,genome size,and ploidy level

Among the traits whose relevance for plant invasions has recently been suggested are genome size (the amount of nuclear DNA) and ploidy level. So far, research on the role of genome size in invasiveness has been mostly based on indirect evidence by comparing species with different genome sizes, but how karyological traits influence competition at the intraspecific level remains unknown. We addressed these questions in a common‐garden experiment evaluating the outcome of direct intraspecific competition among 20 populations of Phragmites australis, represented by clones collected in North America and Europe, and differing in their status (native and invasive), genome size (small and large), and ploidy levels (tetraploid, hexaploid, or octoploid). Each clone was planted in competition with one of the others in all possible combinations with three replicates in 45‐L pots. Upon harvest, the identity of 21 shoots sampled per pot was revealed by flow cytometry and DNA analysis. Differences in performance were examined using relative proportions of shoots of each clone, ratios of their aboveground biomass, and relative yield total (RYT). The performance of the clones in competition primarily depended on the clone status (native vs. invasive). Measured in terms of shoot number or aboveground biomass, the strongest signal observed was that North American native clones always lost in competition to the other two groups. In addition, North American native clones were suppressed by European natives to a similar degree as by North American invasives. North American invasive clones had the largest average shoot biomass, but only by a limited, nonsignificant difference due to genome size. There was no effect of ploidy on competition. Since the North American invaders of European origin are able to outcompete the native North American clones, we suggest that their high competitiveness acts as an important driver in the early stages of their invasion.     

Jack-and-Master Trait Responses to Elevated CO2 and N: A Comparison of Native and Introduced Phragmites australis

Global change is predicted to promote plant invasions world-wide, reducing biodiversity and ecosystem function. Phenotypic plasticity may influence the ability of introduced plant species to invade and dominate extant communities. However, interpreting differences in plasticity can be confounded by phylogenetic differences in morphology and physiology. Here we present a novel case investigating the role of fitness trait values and phenotypic plasticity to global change factors between conspecific lineages of Phragmites australis. We hypothesized that due to observed differences in the competitive success of North American-native and Eurasian-introduced P. australis genotypes, Eurasian-introduced P. australis would exhibit greater fitness in response to global change factors. Plasticity and plant performance to ambient and predicted levels of carbon dioxide and nitrogen pollution were investigated to understand how invasion pressure may change in North America under a realistic global change scenario. We found that the introduced Eurasian genotype expressed greater mean trait values in nearly every ecophysiological trait measured – aboveground and belowground – to elevated CO₂ and nitrogen, outperforming the native North American conspecific by a factor of two to three under every global change scenario. This response is consistent with “jack and master” phenotypic plasticity. We suggest that differences in plant nitrogen productivity, specific leaf area, belowground biomass allocation, and inherently higher relative growth rate are the plant traits that may enhance invasion of Eurasian Phragmites in North America. Given the high degree of genotypic variability within this species, and our limited number of genotypes, our results must be interpreted cautiously. Our study is the first to demonstrate the potential importance of jack-and-master phenotypic plasticity in plant invasions when facing imminent global change conditions. We suggest that jack-and-master invasive genotypes and/or species similar to introduced P. australis will have an increased ecological fitness, facilitating their invasion in both stressful and resource rich environments.     

Neopolyploidy in Spartina pectinata Link: 1. Morphological analysis of tetraploid and hexaploid plants in a mixed natural population

Prairie cordgrass has been reported as a multi-polyploidy species having three cytotypes: tetra- (2n?=?4x?=?40), hexa- (2n?=?6x?=?60), and octoploid (2n?=?8x?=?80). A mixed-ploidy population comprising tetraploids and hexaploids was recently found at a single location in Illinois. However, adaptation and morphological differences between tetra- and hexaploids occurring in natural conditions as well as the contact zones of these cytotypes have yet to be determined. In this study, the cytotypes of 147 individuals of prairie cordgrass collected across the contact zone (4x?+?6x) were determined by flow cytometry using somatic G1 nuclei, and the results were confirmed by chromosome counts. Nineteen morphological characteristics were compared between the cytotypes. Tetra- and hexaploid plants have 2C genome sizes of 1.57 and 2.36?pg with chromosome counts of 40 and 60, respectively. This increase in polyploidy resulted in a greater variability of morphological expression in Illinois prairie cordgrass. Substantial differences in the flowering time, stomatal size, and plant morphological characteristics were observed between tetra- and hexaploids. The results indicate that the increasing of ploidy level in prairie cordgrass resulted in increased plant size in ploidy mixtures. The recent event of ploidy mixtures in prairie cordgrass natural populations offers unique opportunities for studying the formation and establishment of neopolyploidy.     

Contrasting trait responses to latitudinal climate variation in two lineages of an invasive grass

Plants are expected to respond to global environmental change through shifts in functional traits and in their ranges. These shifts could alter productivity and interactions among species or genetic lineages, ultimately leading to changes in distributions and abundance. In particular, cosmopolitan species are predicted to increase growth with decreasing latitude due to differences in climate and temperature. The pattern of changes in growth may vary among genotypes within species, leading to different responses with latitude. To evaluate whether climate can affect geographically distinct genotypes of cosmopolitan invasive species differently, we evaluated the trait responses of two lineages of the common reed, Phragmites australis, to variation in environmental conditions spanning North America’s Atlantic coast. Using three reciprocal transplant common gardens, we tested for the effects of garden location and plant lineage on traits related to biomass production, flowering frequency, leaf morphology, and leaf-level physiology. We found that aboveground biomass, stem density, and flowering frequency responded non-linearly to increasing latitude in one or both lineages. These results suggest that measures of plant traits over narrow latitudinal ranges may not accurately reflect organismal-level responses to global change at broad spatial scales. Given the responses to latitude that we observed in P. australis, we propose that feedbacks between growth and reproductive rate will influence range shifts in these two lineages. Such range shifts could lead to genetic admixtures, subsequently yielding more productive, locally-adapted genotypes.     

Geographic variation in growth responses in Phragmites australis

Phragmites australis is a cosmopolitan wetlands species occurring in a wide range of climatic habitats. It can be assumed that adaptations to climate have evolved to enable the synchronization of growth with the seasonality of the environment. To study these adaptations, European P. australis was collected in different geographic regions, and grown in common environments situated in the Czech Republic, Denmark and The Netherlands.Phragmites australis originating from higher latitudes showed higher relative length growth rates (RLGR), and flowered earlier in time than that from lower latitudes. Plants from Spain even continued growth until the first autumn frosts. When grown in the different common environments, population differences were found in RLGR, but no general trend was apparent. On average, shoots started to grow 2 weeks earlier in The Netherlands than in Denmark and 6 weeks earlier than in the Czech Republic. These differences could be largely related to lower spring temperatures in the latter two countries. When shoot-growth was plotted against the temperature sum, no differences in RLGR between Denmark and The Netherlands were apparent, whereas shoot-growth was slower in the Czech Republic.Results from a greenhouse experiment showed that seedlings from southern populations formed taller but fewer shoots and thicker but shorter rhizomes than those from northern populations, irrespective of total dry weight. They also allocated more dry matter to stems at the expense of leaves, whereas no differences in allocation to below-ground plant parts were found.It was concluded that populations of P. australis showed clinal variation in (i) the length of the growing season, (ii) time of flowering, and (iii) morphology and biomass allocation. These results are discussed with respect to the possible effects of global warming on population functioning.     

Morphology,production and mineral contents inPhragmites australis in different waterbodies of the Estonian SSR

The present work was carried out at four waterbodies in the Estonian SSR where seven different habitats ofPhragmites australis were selected. The paper gives the results of the study of clones from essentially different sites: the characteristics of the habitat of the species (soil profile, soil analyses, pH of water), morphological features of shoots, the nutrient content in plant parts, the production of clones, and also relations between these characteristics.     

Flow cytometric assessment of cytotype distributions within local populations of Phragmites australis (Poaceae) around Lake Biwa,the largest lake in Japan

Phragmites australis (Poaceae) is a clonal perennial that is an important component of wetland ecosystems worldwide. Using flow cytometry, we examined the cytotype distributions within five populations of P. australis located in the vicinity of Lake Biwa. As in previous reports, two ploidy levels, octoploid (2n = 8x = 96) and decaploid (2n = 10x = 120), were identified, which are assumed to be the main cytotypes around the lake. The coexistence of two cytotypes was detected in four of the five populations, suggesting the relatively common occurrence of mixed ploidy levels in the populations around Lake Biwa. Although intermingled cytotype distributions were observed in some populations, the 9x cytotype, that is, the expected outcome of inter‐cytotype crosses, was not observed, indicating limited gene flow between the two cytotypes.     

Trait‐based adaptability of Phragmites australis to the effects of soil water and salinity in the Yellow River Delta

Phragmites australis is the dominant species in the Yellow River Delta and plays an important role in wetland ecosystems. In order to evaluate the relationship between phenotypic variation and environmental factors, explore how functional traits respond to changes in electrical conductivity and soil water content, and reveal the ecological strategies of P. australis, we investigated the ecological responses of P. australis to soil properties based on 96 plots along the coastal–inland regions in the Yellow River Delta of China. Within the range of soil water content (SWC, 9.39%–36.92%) and electrical conductivity (EC, 0.14–13.29 ms/cm), the results showed that (a) the effects of salinity were more important than the soil water content for the characterization of the morphological traits and that plant functional traits including leaf traits and stem traits responded more strongly to soil salinity than soil water content; (b) compared with morphological traits such as average height and internode number, physiological traits such as SPAD value, as well as morphological traits closely related to physiological traits such as specific leaf area and leaf thickness, showed stronger stability in response to soil water and salinity; and (c) under the condition of high electrical conductivity, P. australis improved its water acquisition ability by increasing indicators such as leaf water content and leaf thickness. In addition, with the increase in plant tolerance to stress, more resources were used to resist external stress, and the survival strategy was inclined toward the stress tolerator (S) strategy. Under low EC conditions, P. australis increased specific leaf area and leaf area for its growth in order to obtain resources rapidly, while its survival strategy gradually moved toward the competitor (C) strategy.     

Evidence for multiple introductions of Phragmites australis to North America: detection of a new non-native haplotype

We found a new non-native haplotype of Phragmites australis in North America that provides convincing evidence for multiple introductions of this highly invasive reed from Europe. Prior to our detection of this new non-native haplotype, invasion of North America by this reed grass was thought to be limited to a single cp-DNA haplotype–haplotype M. However, we found two sites colonized by haplotype L1 in Quebec, Canada, a haplotype native to northern Europe, Great Britain and Romania. Because the invasion of North America by P. australis is ongoing, and because there is evidence for intra- and inter-specific hybridization and increased fecundity resulting from outcrossing, more attention should be paid to genetic differences and associated vigor of populations of introduced Phragmites across North America.     

Differences in morphology and C/N-balance between clones ofPhragmites australis within a plantation at a degraded fen

The morphology, productivity and C/N-balance of 9 different clones ofPhragmites australis planted in 1997 in a degraded fen area of 40,000 m² was investigated in order to estimate the degree of variation between the genotypes. The planted reed clones showed significant differences in morphology, standing crop and stand structure at the same site. The above-ground biomass of some reed clones was due to high culms and large leaf areas, while among others it was due to high shoot densities and small culms. The productivity of the individual clones also differed. At the end of the 1998 growing season the standing crop of the clones ranged from about 700 to 2,000 g of dry matter per m². Differences were found in the C/N-dynamics as in the standing stock of total nitrogen in the above-ground biomass (ranging from 15 to 50 g N/m²) and the relative nitrogen content of the shoots. Furthermore, seasonal changes in the amount of free amino acids and carbohydrates in the basal internodes of the different clones were compared. The patterns are discussed with respect to the nutritional status of the reed plants. In conclusion, the results suggest high genotypic variation despite the comparable site conditions and thus a strong influence of genetically determined differences in growth and resource exploitation on the characteristics of reed clones.     

Phragmites australis root secreted phytotoxin undergoes photo-degradation to execute severe phytotoxicity

Our study organism, Phragmites australis (common reed), is a unique invader in that both native and introduced lineages are found coexisting in North America. This allows one to make direct assessments of physiological differences between these different subspecies and examine how this relates to invasiveness. Recent efforts to understand plant invasive behavior show that some invasive plants secrete a phytotoxin to ward-off encroachment by neighboring plants (allelopathy) and thus provide the invaders with a competitive edge in a given habitat. Here we show that a varying climatic factor like ultraviolet (UV) light leads to photo-degradation of secreted phytotoxin (gallic acid) in P. australis rhizosphere inducing higher mortality of susceptible seedlings. The photo-degraded product of gallic acid (hereafter GA), identified as mesoxalic acid (hereafter MOA), triggered a similar cell death cascade in susceptible seedlings as observed previously with GA. Further, we detected the biological concentrations of MOA in the natural stands of exotic and native P. australis. Our studies also show that the UV degradation of GA is facilitated at an alkaline pH, suggesting that the natural habitat of P. australis may facilitate the photo-degradation of GA. The study highlights the persistence of the photo-degraded phytotoxin in the P. australis''s rhizosphere and its inhibitory effects against the native plants.Key words: ultraviolet, gallic acid, mesoxalic acid, novel weapons, invasive species, Phragmites australis     

A phylogeographic study of the cosmopolitan genus Phragmites (Poaceae) based on AFLPs

Within the genus Phragmites (Poaceae), the species P. australis (the common reed) is virtually cosmopolitan, and shows considerable variation in ploidy level and morphology. Genetic variation in Phragmites was studied using AFLPs, and analysed with parsimony and distance methods. Groups of P. australis strongly supported in the analyses include one that comprises all South American clones, a distinct group from the US Gulf Coast, and a group of E. Asian and Australian octoploids. Among the other species, the paleotropical P. vallatoria is supported as monophyletic and most closely related to the paraphyletic P. mauritianus and to the Gulf Coast and S. American groups. The E. Asian species P. japonicus is closely related to a group of P. australis clones mostly from central North America. Tetraploidy predominates in the genus, and optimisation of chromosome numbers onto the phylogeny shows that higher ploidy levels have evolved many times.     

Genetics,novel weapons and rhizospheric microcosmal signaling in the invasion of Phragmites australis

Chemical communication and perception strategies between plants are highly sophisticated but are only partly understood. Among the different interactions, the suppressive interaction of a class of chemicals released by one plant through root exudates against the neighbouring plants (allelopathy) have been implicated in the invasiveness of many exotic weedy species. Phragmites australis (common reed) is one of the dominant colonizers of the North American wetland marshes and exhibits invasive behavior by virtually replacing the entire native vegetation in its niche. Recently, by adopting a systematic bioassay driven approach we elucidated the role of root derived allelopathy as one of the important mechanisms by which P. australis exerts its invasive behavior. Additionally, our recent preliminary data indicates the involvement of rhizobacterial signaling in the invasive success of P. australis. A better understanding of biochemical weaponry used by P. australis will aid scientists and technologists in addressing the impact of root secretions in invasiveness of weedy species and thus promote a more informed environmental stewardship.Key words: Phragmites australis, roots, phytotoxicity, reactive oxygen species, microtubules, microcosm     

Cytogeographic Distribution and Genome Size Variation in Prairie Cordgrass (Spartina pectinata Bosc ex Link)

Prairie cordgrass plants (Spartina pectinata Bosc ex Link) were examined from 61 locations representing the geographic distribution of prairie cordgrass in the U.S. Using flow cytometry, the genome size of 183 individual plants of prairie cordgrass was determined, and the chromosome counts were obtained. Three distinct ploidy levels were observed: tetraploid ( $ \overline x = {1}.{56} {\text{pg}},{2}n = {4} \times = {4}0 $ ), hexaploid ( $ \overline x { = 2}.{33} {\text{pg}},{2}n = {6} \times = {6}0 $ ), and octoploid ( $ \overline {\text{x}} { = 3}.0{6} {\text{pg}},{2}n = {8} \times { = 8}0 $ ). In the sampled areas, the tetraploid populations extended from the East North Central to the New England regions of the U.S., while the octoploid cytotypes were mostly distributed in the West North Central regions. Populations of the tetraploids and octoploids were found in close proximity in the West North Central (Iowa and Kansas) and the West South Central (Oklahoma) regions. The hexaploid cytotype was found in one mixed population (4x?+?6x) occurring in Illinois. No statistically significant intraploidy genome size variation was found in the tetraploid populations, while significant intraploidy genome size variation was found in the octoploid populations. This study precisely defined the geographic distribution of cytotypes in prairie cordgrass throughout the different regions of the U.S. These results provide critical genome size and ploidy distribution information needed to design efficient breeding schemes for high yielding cultivars of prairie cordgrass with local adaption.     

Phytogeographical studies ofCalamagrostis sachalinensis (Gramineae)

Chromosome numbers were determined on 223 collections ofCalamagrostis sachalinensis from 18 localities in Japan. The plants were found to be tetraploid (2n=28), hexaploid (2n=42) or octoploid (2n=56). A few collections were found to include one or two B-chromosomes. The tetraploid collections were made from central Honshu and Mt. Apoi in Hokkaido, while the hexaploids and the octoploids were detected in many localities. Pollen examination of these collections showed that the tetraploids with but one exception have good pollen and the hexaploids and the octoploids have no pollen or have bad pollen with stainability less than 10%. With the help of pollen examination of a number of herbarium specimens, the distribution of the tetraploids and that of the assemblage of the hexaploids and octoploids were delineated. Morphological studies indicated that the tetraploid, hexaploid and octoploid plants can not be separated in gross and spikelet morphology and that the tetraploids in central Honshu and those in Mt. Apoi are significantly different in leaf features. It was concluded thatC. sachalinensis represents an apo-amphimictic complex, which includes the following four races: 1) tetraploid, amphimictic, having thin leaf blades 5–10 mm broad and growing on the subalpine conifer forest belt and the conifer forest-alpine ecotone in the mountains of central Honshu; 2) tetraploid, amphimictic, having hard leaf blades 2–6 mm broad and growing on the stony, arid and exposed alpine belt on Mt. Apoi in Hokkaido; 3) hexaploid, mainly apomictic, the most variable ecologically, widely distributed; 4) octoploid, mainly apomictic, frequent in the upper montane to alpine belts, probably widely distributed.     

A morphological and chemical analysis of geographical variation in Tilia L. of eastern North America

A statistical analysis of morphological variation and a Chromatographic analysis of flavonoid variation were performed to determine taxonomic relationships among the species ofTilia of eastern North America. No apparent morphological discontinuities were seen between populations within the sample area although two characters (involving leaf pubescence and gland length) showed definite patterns of geographical variation. Flavonoid patterns showed definite differences between northern and southern populations with an intermediate zone in the Smoky Mountain region. The continuous nature of the morphological and flavonoid variation suggested that the genus as represented in eastern North America should be regarded as one species,Tilia americana L.     

Phenotypic plasticity of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> and <Emphasis Type="Italic">Phragmites australis</Emphasis> in response to nitrogen addition and intraspecific competition

Phenotypic plasticity of the two salt marsh grasses Spartina alterniflora and Phragmites australis in salt marshes is crucial to their invasive ability, but the importance of phenotypic plasticity, nitrogen levels, and intraspecific competition to the success of the two species is unclear at present. Spartina alterniflora Loisel. is an extensively invasive species that has increased dramatically in distribution and abundance on the Chinese and European coasts, and has had considerable ecological impacts in the regions where it has established. Meanwhile, Phragmites australis Cav., a native salt marsh species on the east coast of China, has replaced the native S. alterniflora in many marshes along the Atlantic Coast of the US. This study determined the effects of nitrogen availability and culm density on the morphology, growth, and biomass allocation traits of Spartina alterniflora and Phragmites australis. A large number of morphological, growth, and biomass parameters were measured, and various derived values (culm: root ratio, specific leaf area, etc.) were calculated, along with an index of phenotypic plasticity. Nitrogen addition significantly affected growth performance and biomass allocation traits of Spartina alterniflora, and culm density significantly affected morphological characteristics in a negative way, especially for Spartina alterniflora. However, there were no significant interactions between nitrogen levels and culm density on the morphological parameters, growth performances parameters, and biomass allocation parameters of the two species. Spartina alterniflora appears to respond more strongly to nitrogen than to culm density and this pattern of phenotypic plasticity appears to offer an expedition for successful invasion and displacement of Phramites australias in China. The implication of this study is that, in response to the environmental changes that are increasing nitrogen levels, the range of Spartina alterniflora is expected to continue to expand on the east coast of China.     

Meiotic pairing as an indicator of genome composition in polyploid prairie cordgrass (<Emphasis Type="Italic">Spartina pectinata</Emphasis> Link)

The existence of neopolyploidy in prairie cordgrass (Spartina pectinata Link) has been documented. The neohexaploid was discovered coexisting with tetraploids in central Illinois, and has been reported to exhibit competitiveness in the natural environment. It is hypothesized that the natural tetraploid cytotype produced the hexaploid cytotype via production of unreduced gametes. Meiosis I chromosome pairing was observed in tetraploid (2n?=?4x?=?40), hexaploid (2n?=?6x?=?60), and octoploid (2n?=?8x?=?80) accessions and the percentage of meiotic abnormality was determined. Significant differences in meiotic abnormality exist between tetraploid, hexaploid, and octoploid cytotypes. An elevated incidence of abnormal, predominantly trivalent pairing in the neohexaploid suggests that it may possess homologous chromosomes in sets of three, in contrast to the tetraploid and octoploid cytotypes, which likely possess homologous chromosomes in sets of two. Abnormal chromosome pairing in the hexaploid may result in unequal allocation of chromosomes to daughter cells during later stages of meiosis. Chromosome pairing patterns in tetraploid, hexaploid, and octoploid cytotypes indicate genome compositions of AABB, AAABBB, and AABBA′A′B′B′, respectively.     

A new cytotype of Jacobaea vulgaris (Asteraceae): frequency,morphology and origin

Jacobaea vulgaris subsp. vulgaris (syn. Senecio jacobaea subsp. jacobaea) constitutes an intricate polyploid complex distributed in Europe. Four cytotypes have been reported in this species, three with euploid (diploid, tetraploid and octoploid; 2n=20, 40 and 80) and one with aneuploid (2n=32) chromosome numbers. Here we report that the diploid chromosome number (2n=20) reported from Bulgaria is due to misidentification with Jacobaea aquatica. On the other hand, we have discovered a new, hexaploid (2n=6x=60) cytotype within J. vulgaris subsp. vulgaris using flow cytometry. The new cytotype occurs within four sympatric populations of otherwise tetraploid and octoploid plants in Pannonia (one locality in the eastern Czech Republic and two localities in southwestern Slovakia) and in Podillya (one locality in western Ukraine). The frequency of hexaploid individuals within 76 studied populations is very low (only 10 of 693 analysed plants), and hexaploids probably represent hybrids between tetraploid and octoploid plants. Three mixed populations with hexaploid plants were subjected to detailed morphological and pollen fertility analyses. Multivariate morphometric analysis reveals partial separation of tetraploid and octoploid plants, whereas hexaploid individuals are similar in morphology to octoploids. In comparison with tetraploids, octoploids and hexaploids exhibit slightly longer ray florets, involucral bracts and tubular florets and more hairy outer achenes. Hexaploid plants display larger pollen grains and lower pollen fertility compared to tetraploids and octoploids.