Here we describe the results of a regional comparison of introduced Phragmites
australis and two other P. australis lineages found in North
America. The regional similarities and differences in introduced P.
australis invasion highlight the importance of continental-scale studies for
decoding plant invasions.
We use a regional comparison of Phragmites australis (common reed)
subsp. americanus, P. australis subsp.
berlandieri and introduced P. australis (possibly
five sublineages) in the Chesapeake Bay, the St Lawrence River, Utah and the Gulf Coast
to inform a North American perspective on P. australis invasion
patterns, drivers, impacts and research needs.
Findings and research needs
Our regional assessments reveal substantial diversity within and between the three main
lineages of P. australis in terms of mode of reproduction and the types
of environment occupied. For introduced P. australis, the timing of
introduction also differed between the regions. Nevertheless, a common finding in these
regions reinforces the notion that introduced P. australis is
opportunistic and thrives in disturbed habitats. Thus, we expect to see substantial
expansion of introduced P. australis with increasing anthropogenic
disturbances in each of these regions. Although there have been some studies documenting
the negative impacts of introduced P. australis, it also plays a
beneficial role in some regions, and in some cases, the purported negative impacts are
unproven. There is also a broader need to clarify the genetic and ecological
relationships between the different introduced sublineages observed in North America,
and their relative competitive ability and potential for admixture. This may be done
through regional studies that use similar methodologies and share results to uncover
common patterns and processes. To our knowledge, such studies have not been performed on
P. australis in spite of the broad attention given to this species.
Such research could advance theoretical knowledge on biological invasion by helping to
determine the extent to which the patterns observed can be generalized or are sublineage
specific or region specific.
Given what appears to be sometimes idiosyncratic invasion patterns when interpreted in
isolation in the regions that we analysed, it may be time to consider initiatives on a
continental (if not intercontinental) scale to tackle unresolved issues about P.
European Phragmites australis is one of four main cp-DNA haplotype clusters present worldwide. The European gene pool extends from North America to Far East Asia and South Africa. Extensive gene flow occurs only within the temperate region of Europe.
Background and aims
Two Phragmites australis taxa are recognized in Europe: P. australis ssp. altissimus, also known as Phragmites isiaca, in the Mediterranean region and P. australis in the temperate region. Another taxonomic group in the Mediterranean is Phragmites frutescens. European genotypes are diverse genetically, cytologically and morphologically, and are related to African, Asiatic and American genotypes. We investigated chloroplast DNA (cpDNA) diversity in Europe and defined the current borders of the European gene pool.
We analysed chloroplast variation with parsimony and genetic distance methods, and compared it with that of nuclear amplified fragment length polymorphism and microsatellites. We also investigated the phenological pattern of 188 genotypes collected worldwide in a common garden in Denmark. We assumed that non-flowering genotypes could indicate climatic, geographic and/or reproductive barriers to dispersal and would have been recorded in the genetic pattern as groups genetically isolated from, or within, the European pool.
The European P. australis gene pool extends from North America to the Far East and South Africa. However, African and North American genotypes are differentiating from the European genotypes. Mediterranean P. australis is genetically different from temperate P. australis and shares several similarities with Phragmites mauritianus in Africa and Phragmites karka in Asia. Phragmites frutescens shares the cpDNA sequences with both these tropical species. Two DNA bands can distinguish Mediterranean P. australis from P. frutescens and P. mauritianus and from temperate P. australis, and reveal possible hybrids among these species in the Mediterranean region. Phenological data confirmed possible gene flow within the temperate region of Europe, whereas the Mediterranean genotypes did not set inflorescences in Denmark, suggesting reproductive barriers between temperate and Mediterranean P. australis.
European P. australis appears as one of four main Phragmites groups known in the world. Further research is needed to understand the implications of long-distance dispersal at the population level.
Nonnative Phragmites australis (common reed) is one of the most intensively researched and managed invasive plant species in the United States, yet as with many invasive species, our ability to predict, control or understand the consequences of invasions is limited. Rapid spread of dense Phragmites monocultures has prompted efforts to limit its expansion and remove existing stands. Motivation for large-scale Phragmites eradication programs includes purported negative impacts on native wildlife, a view based primarily on observational results. We took an experimental approach to test this assumption, estimating the effects of nonnative Phragmites australis on a native amphibian.
Concurrent common garden and reciprocal transplant field experiments revealed consistently strong positive influences of Phragmites on Rana catesbeiana (North American bullfrog) larval performance. Decomposing Phragmites litter appears to contribute to the effect.
Positive effects of Phragmites merit further research, particularly in regions where both Phragmites and R. catesbeiana are invasive. More broadly, the findings of this study reinforce the importance of experimental evaluations of the effects of biological invasion to make informed conservation and restoration decisions.
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.
ultraviolet; gallic acid; mesoxalic acid; novel weapons; invasive species; 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.
Phragmites australis; roots; phytotoxicity; reactive oxygen species; microtubules; microcosm
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.
The interactions among Spartina patens and sediment microbial populations and the interactions among Phragmites australis and sediment microbial populations were studied at monotypic sites in Piermont Marsh, a salt marsh of the Hudson River north of New York, N.Y., at key times during the growing season. Arbuscular mycorrhizal fungi (AMF) effectively colonized S. patens but not P. australis, and there were seasonal increases and decreases that coincided with plant growth and senescence (17 and 6% of the S. patens root length were colonized, respectively). In sediment samples from the Spartina site, the microbial community and specific bacterial populations were at least twice as large in terms of number and biomass as the microbial community and specific bacterial populations in sediment samples from the Phragmites site, and peak values occurred during reproduction. Members of the domain Bacteria, especially members of the α-, γ-, and δ-subdivisions of the Proteobacteria, were the most abundant organisms at both sites throughout the growing season. The populations were generally more dynamic in samples from the Spartina site than in samples from the Phragmites site. No differences between the two sites and no differences during the growing season were observed when restriction fragment length polymorphism analyses of nifH amplicons were performed in an attempt to detect shifts in the diversity of nitrogen-fixing bacteria. Differences were observed only in the patterns generated by PCR or reverse transcription-PCR for samples from the Spartina site, suggesting that there were differences in the overall and active populations of nitrogen-fixing bacteria. Regression analyses indicated that there was a positive interaction between members of the δ-subdivision of the Proteobacteria and root biomass but not between members of the δ-subdivision of the Proteobacteria and macroorganic matter at both sites. In samples from the Spartina site, there were indications that there were bacterium-fungus interactions since populations of members of the α-subdivision of the Proteobacteria were negatively associated with AMF colonization and populations of members of the γ-subdivision of the Proteobacteria were positively associated with AMF colonization.
The invasive species Spartina alterniora Loisel was introduced to the eastern coast of China in the 1970s and 1980s for the purposes of land reclamation and the prevention of soil erosion. The resulting interspecific competition had an important influence on the distribution of native vegetation, which makes studying the patterns and mechanisms of the interactions between Spartina alterniora Loisel and the native species Phragmites australis (Cav.) Trin ex Steud in this region very important. There have been some researches on the interspecific interactions between P. australis and S. alterniora in the Dongtan wetland of Chongming, east China, most of which has focused on the comparison of their physiological characteristics. In this paper, we conducted a neighbor removal experiment along a tidal gradient to evaluate the relative competitive abilities of the two species by calculating their relative neighbor effect (RNE) index. We also looked at the influence of environmental stress and disturbance on the competitive abilities of the two species by comparing interaction strength (I) among different tidal zones both for P. australis and S. alterniora. Finally, we measured physiological characteristics of the two species to assess the physiological mechanisms behind their different competitive abilities. Both negative and positive interactions were found between P. australis and S. alterniora along the environmental gradient. When the direction of the competitive intensity index for P. australis and S. alterniora was consistent, the competitive or facilitative effect of S. alterniora on P. australis was stronger than that of P. australis on S. alterniora. The interspecific interactions of P. australis and S. alterniora varied with environmental conditions, as well as with the method used, to measure interspecific interactions.
The present study reveals significant genetically determined differences in a range of growth and ecophysiological traits between different Phragmites australis genotypes, and provides evidence that the differences are neither related to ploidy level per se nor to the phylogeographic relationships of the genotypes.
Background and aims
Phragmites australis is a wetland grass with high genetic variability, augmented by its cosmopolitan distribution, clonal growth form and large variation in chromosome numbers. Different ploidy levels and ecotypes differ in morphology and ecophysiological traits, and may possess different levels of phenotypic variation. The aim of this study was to quantify the natural variation in ecophysiological characteristics of P. australis, and to explore whether differences in ecophysiological traits can be related to ploidy levels or to the geographic origin of the clones.
Fifteen clones of P. australis from Europe and Asia/Australia, representing five ploidy levels (4x, 6x, 8x, 10x and 12x), were grown in a common garden design for 119 days. Plant growth and light-saturated rate of photosynthesis (Pmax), stomatal conductance (gs), water use efficiency (WUE) and concentrations of photosynthetic pigments and mineral ions in the leaves were measured.
The growth of the plants and most ecophysiological parameters differed significantly between clones. The mean maximum shoot height varied from 0.9 to 1.86 m, Pmax from 9.7 to 27 µmol m−2 s−1, gs from 0.22 to 1.41 mol m−2 s−1 and WUE from 13 to 47 µmol mol−1. The concentrations of chlorophylls did not vary significantly between clones, but the chlorophyll a/b ratio and the concentrations of total carotenoids did. The observed differences were not explained either by the ploidy level per se or by the geographic origin or phylogenetic relationships of the clones.
Phylogeographic relationships in P. australis on a global scale do not mirror the environment where the adaptations have evolved, and high phenotypic variation among and within clones complicates comparative studies. Future studies aimed at explaining differences in plant behaviour between P. australis populations should be careful in the selection of target genotypes and/or populations, and should avoid generalizing their findings beyond the genotypes and/or populations studied.
Assessing oil pollution using traditional field-based methods over large areas is difficult and expensive. Remote sensing technologies with good spatial and temporal coverage might provide an alternative for monitoring oil pollution by recording the spectral signals of plants growing in polluted soils. Total petroleum hydrocarbon concentrations of soils and the hyperspectral canopy reflectance were measured in wetlands dominated by reeds (Phragmites australis) around oil wells that have been producing oil for approximately 10 years in the Yellow River Delta, eastern China to evaluate the potential of vegetation indices and red edge parameters to estimate soil oil pollution. The detrimental effect of oil pollution on reed communities was confirmed by the evidence that the aboveground biomass decreased from 1076.5 g m−2 to 5.3 g m−2 with increasing total petroleum hydrocarbon concentrations ranging from 9.45 mg kg−1 to 652 mg kg−1. The modified chlorophyll absorption ratio index (MCARI) best estimated soil TPH concentration among 20 vegetation indices. The linear model involving MCARI had the highest coefficient of determination (R2 = 0.73) and accuracy of prediction (RMSE = 104.2 mg kg−1). For other vegetation indices and red edge parameters, the R2 and RMSE values ranged from 0.64 to 0.71 and from 120.2 mg kg−1 to 106.8 mg kg−1 respectively. The traditional broadband normalized difference vegetation index (NDVI), one of the broadband multispectral vegetation indices (BMVIs), produced a prediction (R2 = 0.70 and RMSE = 110.1 mg kg−1) similar to that of MCARI. These results corroborated the potential of remote sensing for assessing soil oil pollution in large areas. Traditional BMVIs are still of great value in monitoring soil oil pollution when hyperspectral data are unavailable.
Roads, bridges, and dikes constructed across salt marshes can restrict tidal flow, degrade habitat quality for nekton, and facilitate invasion by non-native plants including Phragmites australis. Introduced P. australis contributes to marsh accretion and eliminates marsh surface pools thereby adversely affecting fish by reducing access to intertidal habitats essential for feeding, reproduction, and refuge. Our study assessed the condition of resident fish populations (Fundulus heteroclitus) at four tidally restricted and four tidally restored marshes in New England invaded by P. australis relative to adjacent reference salt marshes. We used physiological and morphological indicators of fish condition, including proximate body composition (% lipid, % lean dry, % water), recent daily growth rate, age class distributions, parasite prevalence, female gravidity status, length-weight regressions, and a common morphological indicator (Fulton’s K) to assess impacts to fish health. We detected a significant increase in the quantity of parasites infecting fish in tidally restricted marshes but not in those where tidal flow was restored to reduce P. australis cover. Using fish length as a covariate, we found that unparasitized, non-gravid F. heteroclitus in tidally restricted marshes had significantly reduced lipid reserves and increased lean dry (structural) mass relative to fish residing in reference marshes. Fish in tidally restored marshes were equivalent across all metrics relative to those in reference marshes indicating that habitat quality was restored via increased tidal flushing. Reference marshes adjacent to tidally restored sites contained the highest abundance of young fish (ages 0–1) while tidally restricted marshes contained the lowest. Results indicate that F. heteroclitus residing in physically and hydrologically altered marshes are at a disadvantage relative to fish in reference marshes but the effects can be reversed through ecological restoration.
Interbreeding potential, chromosome number, and host range were compared among several isolates and species of Bursaphelenchus from diverse geographic areas. Some isolates from North America, Japan, and France had a wide-ranging interbreeding potential, whereas others were restricted in their potential to hybridize with other isolates. Although interbreeding occurred in the laboratory between some "M" and "R" forms of B. xylophilus, interbreeding of B. xylophilus and B. mucronatus was rare. The hybrids had the pathogenicity of the parent with the broader host range. This fact suggests that virulence may be inherited as a dominant character or that increased virulence may have resulted from differences in hybrid vigor. The haploid chromosome number of the different isolates separated the isolates into three groups and distinguished B. xylophilus from B. mucronatus. The findings suggest that the pinewood nematode species complex consists of sibling species that have evolved by reproductive isolation, that the French isolate is a new species, and that B. xylophilus and B. mucronatus have evolved from a common ancestor.
Bursaphelenchus; chromosome; interbreeding; nematode; pinewood nematode; Pinus; speciation
Contemporary evolution through human-induced hybridization occurs throughout the taxonomic range. Formerly allopatric species appear especially susceptible to hybridization. Consequently, hybridization is expected to be more common in regions with recent sympatry owing to human activity than in areas of historical range overlap. Coyotes (Canis latrans) and gray wolves (C. lupus) are historically sympatric in western North America. Following European settlement gray wolf range contracted, whereas coyote range expanded to include eastern North America. Furthermore, wolves with New World (NW) mitochondrial DNA (mtDNA) haplotypes now extend from Manitoba to Québec in Canada and hybridize with gray wolves and coyotes. Using mtDNA and 12 microsatellite markers, we evaluated levels of wolf-coyote hybridization in regions where coyotes were present (the Canadian Prairies, n = 109 samples) and absent historically (Québec, n = 154). Wolves with NW mtDNA extended from central Saskatchewan (51°N, 69°W) to northeastern Québec (54°N, 108°W). On the Prairies, 6.3% of coyotes and 9.2% of wolves had genetic profiles suggesting wolf-coyote hybridization. In contrast, 12.6% of coyotes and 37.4% of wolves in Québec had profiles indicating hybrid origin. Wolves with NW and Old World (C. lupus) mtDNA appear to form integrated populations in both regions. Our results suggest that hybridization is more frequent in historically allopatric populations. Range shifts, now expected across taxa following climate change and other human influence on the environment, might therefore promote contemporary evolution by hybridization.
Canis; coyote; hybridization; sympatry; wolf
The ability to express phenotypically plastic responses to environmental cues might be adaptive in changing environments. We studied phenotypic plasticity in mating behaviour as a response to population density and adult sex ratio in a freshwater isopod (Asellus aquaticus). A. aquaticus has recently diverged into two distinct ecotypes, inhabiting different lake habitats (reed Phragmites australis and stonewort Chara tomentosa, respectively). In field surveys, we found that these habitats differ markedly in isopod population densities and adult sex ratios. These spatially and temporally demographic differences are likely to affect mating behaviour. We performed behavioural experiments using animals from both the ancestral ecotype (“reed” isopods) and from the novel ecotype (“stonewort” isopods) population. We found that neither ecotype adjusted their behaviour in response to population density. However, the reed ecotype had a higher intrinsic mating propensity across densities. In contrast to the effects of density, we found ecotype differences in plasticity in response to sex ratio. The stonewort ecotype show pronounced phenotypic plasticity in mating propensity to adult sex ratio, whereas the reed ecotype showed a more canalised behaviour with respect to this demographic factor. We suggest that the lower overall mating propensity and the phenotypic plasticity in response to sex ratio have evolved in the novel stonewort ecotype following invasion of the novel habitat. Plasticity in mating behaviour may in turn have effects on the direction and intensity of sexual selection in the stonewort habitat, which may fuel further ecotype divergence.
A method that can be used to measure the initial decomposition rates of polysaccharides in sediment samples was developed. It uses toluene to specifically inhibit microbial uptake of carbohydrates without affecting extracellular hydrolysis of polysaccharides. Accumulating carbohydrates were determined by high-performance liquid chromatography. Field-sampled litter from the common reed (Phragmites australis), which contains cellulose and arabinoxylan as its main polysaccharides, was used as a model system. Toluene concentrations of between 1 and 10% resulted in the accumulation of similar amounts of monomeric carbohydrates, which was linear over time for most neutral sugars. Toluene (3%) did not have an effect on extracellular enzyme activities, and microbial sugar uptake was completely inhibited, as demonstrated with (sup14)C-labelled xylose and glucose. Experiments with enhancement cultures and fixed reed litter suggested that enzymatic hydrolysis of polysaccharides in reed litter was the main source of glucose, xylose, arabinose, and galactose accumulation. In contrast, the accumulation of high amounts of the alditols mannitol and glucitol was probably caused by lysis of the microbial population in toluene-treated reed litter. Glucose accumulated at rates of 1.3 and 0.10 (mu)mol (middot) g of dry matter content(sup-1) (middot) h(sup-1) under aerobic and anaerobic conditions, respectively, whereas xylose accumulation rates were only 10% of the glucose accumulation rates.
The role of hybridization in generating diversity in animals is an active area of discovery and debate. We assess hybridization across a contact zone of northern (Myodes rutilus) and southern (M. gapperi) red-backed voles using variation in skeletal features and both mitochondrial and nuclear loci. This transect extends approximately 550 km along the North Pacific Coast of North America and encompasses 26 populations (n = 485). We establish the history, geographic extent and directionality of hybridization, determine whether hybridization is ongoing, and assess the evolutionary stability of novel genomic combinations.
Identification of M. rutilus and M. gapperi based on the degree of closure of the post-palatal bridge was concordant with the distribution of diagnostic nuclear MYH6 alleles; however, an 80 km zone of introgressed populations was identified. The introgressant form is characterized by having mitochondrial haplotypes closely related to the northern M. rutilus on a nuclear background and morphological characteristics of southern M. gapperi.
Introgression appears to have been historic as pure populations of M. rutilus are now isolated to the north from introgressants or pure M. gapperi by the LeConte Glacier. As we do not find pure M. rutilus or M. gapperi individuals throughout the distribution of the introgressant form, it appears that the introgressants are a self-sustaining entity not requiring continued hybridization between pure parental forms to generate this novel combination of characters.
This study evaluated biodegradation of the insecticide deltamethrin (1 μg l−1) by pure cultures of neustonic (n = 25) and epiphytic (n = 25) bacteria and by mixed cultures (n = 1), which consisted of a mixture of 25 bacterial strains isolated from the surface microlayer (SM ≈ 250 μm) and epidermis of the Common Reed (Phragmites australis, (Cav.) Trin. ex Steud.) growing in the littoral zone of eutrophic lake Chełmżyńskie. Results indicate that neustonic and epiphytic bacteria are characterized by a similar average capacity to degrade deltamethrin. After a 15-day incubation, bacteria isolated from the surface microlayer reduced the initial concentration of deltamethrin by 60%, while the average effectiveness of the bacteria found on the Common Reed equaled 47%.
Biodegradation; Deltamethrin; Epiphytic bacteria; Neustonic bacteria; Lake; Pesticides
Determination of spatial genetic structure (SGS) in natural populations is important for both theoretical aspects of evolutionary genetics and their application in species conservation and ecological restoration. In this study, we examined genetic diversity within and among the natural populations of a cosmopolitan grass Phragmites australis (common reed) in the Yellow River Delta (YRD), China, where a mosaic of habitat patches varying in soil salinity was detected. We demonstrated that, despite their close geographic proximity, the common reed populations in the YRD significantly diverged at six microsatellite loci, exhibiting a strong association of genetic variation with habitat heterogeneity. Genetic distances among populations were best explained as a function of environmental difference, rather than geographical distance. Although the level of genetic divergence among populations was relatively low (F’ST = 0.073), weak but significant genetic differentiation, as well as the concordance between ecological and genetic landscapes, suggests spatial structuring of genotypes in relation to patchy habitats. These findings not only provided insights into the population dynamics of common reed in changing environments, but also demonstrated the feasibility of using habitat patches in a mosaic landscape as test systems to identify appropriate genetic sources for ecological restoration.
Plants are naturally colonized by many fungal species that produce effects ranging from beneficial to pathogenic. However, how many of these fungi are linked with a single host plant has not been determined. Furthermore, the composition of plant-associated fungal communities has not been rigorously determined. We investigated these essential issues by employing the perennial wetland reed Phragmites australis as a model. DNA extracted from roots, rhizomes, stems, and leaves was used for amplification and cloning of internal transcribed spacer rRNA gene fragments originating from reed-associated fungi. A total of 1,991 clones from 15 clone libraries were differentiated by restriction fragment length polymorphism analyses into 345 operational taxonomical units (OTUs). Nonparametric estimators for total richness (Chao1 and ACE) and also a parametric log normal model predicted a total of about 750 OTUs if the libraries were infinite. Sixty-two percent of the OTUs sequenced were novel at a threshold of 3%. Several of these OTUs represented undocumented fungal species, which also included higher taxonomic levels. In spite of the high diversity of the OTUs, the mycofloras of vegetative organs were dominated by just a few typical fungi, which suggested that competition and niche differentiation influence the composition of plant-associated fungal communities. This suggestion was independently supported by the results of nested PCR assays specifically monitoring two OTUs over 3 years, which revealed significant preferences for host habitat and host organ.
Fungal endophyte communities are often comprised of many species colonizing the same host. However, little is known about the causes of this diversity. On the one hand, the apparent coexistence of closely related species may be explained by the traditional niche differentiation hypothesis, which suggests that abiotic and/or biotic factors mediate partitioning. For endophytes, such factors are difficult to identify, and are therefore in most cases unknown. On the other hand, there is the neutral hypothesis, which suggests that stochastic factors may explain high species diversity. There is a need to investigate to what extent each of these hypotheses may apply to endophytes.
The niche partitioning of two closely related fungal endophytes, Microdochium bolleyi and M. phragmitis, colonizing Phragmites australis, was investigated. The occurrences of each species were assessed using specific nested-PCR assays for 251 field samples of common reed from Lake Constance, Germany. These analyses revealed niche preferences for both fungi. From three niche factors assessed, i.e. host habitat, host organ and season, host habitat significantly differentiated the two species. M. bolleyi preferred dry habitats, whereas M. phragmitis prevailed in flooded habitats. In contrast, both species exhibited a significant preference for the same host organ, i.e. roots. Likewise the third factor, season, did not significantly distinguish the two species. Differences in carbon utilization and growth temperature could not conclusively explain the niches. The inclusion of three unrelated species of Ascomycota, which also colonize P. australis at the same locations, indicated spatio-temporal niche partitioning between all fungi. None of the species exhibited the same preferences for all three factors, i.e. host habitat, host organ, and time of the season.
The fungal species colonizing common reed investigated in this study seem to exploit niche differences leading to a separation in space and time, which may allow for their coexistence on the same host. A purely neutral model is unlikely to explain the coexistence of closely related endophytes on common reed.
Herbaspirillum sp. strain GW103 was isolated from rhizosphere soil of the reed Phragmites australis on reclaimed land. Here we report the 5.05-Mb draft genome sequence of the strain, providing bioinformation about the agronomic benefits of this strain, such as multiple traits relevant to plant root colonization and plant growth promotion.
A high-affinity K+ transporter PutHKT2;1 cDNA was isolated from the salt-tolerant plant Puccinellia tenuiflora. Expression of PutHKT2;1 was induced by both 300 mM NaCl and K+-starvation stress in roots, but only slightly regulated by those stresses in shoots. PutHKT2;1 transcript levels in 300 mM NaCl were doubled by the depletion of potassium. Yeast transformed with PutHKT2;1, like those transformed with PhaHKT2;1 from salt-tolerant reed plants (Phragmites australis), (i) were able to take up K+ in low K+ concentration medium or in the presence of NaCl, and (ii) were permeable to Na+. This suggests that PutHKT2;1 has a high affinity K+-Na+ symport function in yeast. Arabidopsis over-expressing PutHKT2;1 showed increased sensitivities to Na+, K+, and Li+, while Arabidopsis over-expressing OsHKT2;1 from rice (Oryza sativa) showed increased sensitivity only to Na+. In contrast to OsHKT2;1, which functions in Na+-uptake at low external K+ concentrations, PutHKT2;1 functions in Na+-uptake at higher external K+ concentrations. These results show that the modes of action of PutHKT2;1 in transgenic yeast and Arabidopsis differ from the mode of action of the closely related OsHKT2;1 transporter.
Potassium transporter; Puccinellia tenuiflora; PutHKT2;1; salt stress; salt tolerance
Soil disturbance has been widely recognized as an important factor influencing the structure and dynamics of plant communities. Although soil reworkers were shown to increase habitat complexity and raise the risk of plant invasion, their role in regulating the interactions between native and invasive species remains unclear. We proposed that crab activities, via improving soil nitrogen availability, may indirectly affect the interactions between invasive Spartina alterniflora and native Phragmites australis and Scirpus mariqueter in salt marsh ecosystems. We conducted a two-year mesocosm experiment consisting of five species combinations, i.e., monocultures of three species and pair-wise mixtures of invasive and native species, with crabs being either present or absent for each combination. We found that crabs could mitigate soil nitrogen depletion in the mesocosm over the two years. Plant performance of all species, at both the ramet-level (height and biomass per ramet) and plot-level (density, total above- and belowground biomass), were promoted by crab activities. These plants responded to crab disturbance primarily by clonal propagation, as plot-level performance was more sensitive to crabs than ramet-level. Moreover, crab activities altered the competition between Spartina and native plants in favor of the former, since Spartina was more promoted than native plants by crab activities. Our results suggested that crab activities may increase the competition ability of Spartina over native Phragmites and Scirpus through alleviating soil nitrogen limitation.
Spartina alterniflora has widely invaded the saltmarshes of the Yangtze River Estuary and brought negative effects to the ecosystem. Remote sensing technique has recently been used to monitor its distribution, but the similar morphology and canopy structure among S. alterniflora and its neighbor species make it difficult even with high-resolution images. Nevertheless, these species have divergence on phenological stages throughout the year, which cause distinguishing spectral characteristics among them and provide opportunities for discrimination. The field spectra of the S. alterniflora community as well as its major victims, native Phragmites australis and Scirpus mariqueter, were measured in 2009 and 2010 at multi-phenological stages in the Yangtze River Estuary, aiming to find the most appropriate periods for mapping S. alterniflora. Collected spectral data were analyzed separately for every stage firstly by re-sampling reflectance curves into continued 5-nm-wide hyper-spectral bands and then by re-sampling into broad multi-spectral bands – the same as the band ranges of the TM sensor, as well as calculating commonly used vegetation indices. The results showed that differences among saltmarsh communities’ spectral characteristics were affected by their phenological stages. The germination and early vegetative growth stage and the flowering stage were probably the best timings to identify S. alterniflora. Vegetation indices like NDVI, ANVI, VNVI, and RVI are likely to enhance spectral separability and also make it possible to discriminate S. alterniflora at its withering stage.
Hybridization has been hypothesized to influence invasion through the generation of novel phenotypes and/or increased levels of genetic variance. Based on morphology, hybrids between diffuse knapweed and spotted knapweed, two invasive plants in North America, are present in the invaded range. Some individuals within most diffuse knapweed sites in North America exhibit intermediate diffuse × spotted floral morphology. We examined hybridization at the molecular level, using amplified fragment length polymorphisms. Approximately a quarter of the assayed North American diffuse knapweed individuals exhibited evidence of introgression from spotted knapweed. However, plants with intermediate morphology did not show evidence of mixed ancestry more often than the plants with typical diffuse knapweed morphology. The high proportion of hybrid individuals in North American diffuse knapweed sites found here, combined with evidence from recent studies, suggests that diffuse knapweed was likely introduced with admixed individuals, and the hybrids are not newly created postintroduction. A century of backcrossing with diffuse knapweed has likely decoupled the relationship between morphology and admixture at the molecular level. In contrast to the scenario encountered in North America, in the native range where diploid diffuse and spotted knapweed overlap, hybrid swarms are common. In such sites, the floral phenotype aligns more closely with the genotype.
amplified fragment length polymorphism; biological invasion; diffuse knapweed; hybridization; spotted knapweed; STRUCTURE