Nanocomposite Hydrogels Based on Poly(vinyl alcohol) and Carbon Nanotubes for NIR-Light Triggered Drug Delivery.

Photothermal nanocomposite hydrogels are promising materials for remotely triggering drug delivery by near-infrared (NIR) radiation stimuli. In this work, a novel hydrogel based on poly(vinyl alcohol), poly(vinyl methyl ether-alt-maleic acid), poly(vinyl methyl ether), and functionalized multiwalled carbon nanotubes (MWCNT-f) was prepared by the freeze/thaw method. A comparative characterization of materials (with and without MWCNT-f) was carried out by infrared spectroscopy, differential scanning calorimetry, scanning electron microscopy, mechanical assays, swelling kinetics measurements, and photothermal analysis under NIR irradiation. Hydrophilic chemotherapeutic 5-fluorouracil (5-FU) and hydrophobic ibuprofen drugs were independently loaded into hydrogels, and the drug release profiles were obtained under passive and NIR-irradiation conditions. The concentration-dependent cytotoxicity of materials was studied in vitro using noncancerous cells and cancer cells. Notable changes in the microstructure and physicochemical properties of hydrogels were observed by adding a low content (0.2 wt %) of MWCNT-f. The cumulative release amounts of 5-FU and ibuprofen from the hydrogel containing MWCNT-f were significantly increased by 21 and 39%, respectively, through the application of short-term NIR irradiation pulses. Appropriate concentrations of the nanocomposite hydrogel loaded with 5-FU produced cytotoxicity in cancer cells without affecting noncancerous cells. The overall properties of the MWCNT-f-containing hydrogel and its photothermal behavior make it an attractive material to promote the release of hydrophilic and hydrophobic drugs, depending on the treatment requirements.

Wrack Burial Limits Germination and Establishment of Yellow Flag Iris (Iris pseudacorus L.).

Seed burial under wrack, mats of water-transported plant debris, can limit recruitment of seedlings in wetlands. In a greenhouse experiment, we studied the effects of wrack burial (0, 1, 2, 4, 8 cm depths) on germination and emergence of the macrophyte Iris pseudacorus, native to Europe, Mediterranean Basin, and western Asia, that has invaded wetlands in nearly every global ecozone. We recorded the percentages of germinating, senescent, and quiescent seeds and evaluated seedling establishment and growth relative to substrate environmental variables. Seedling emergence of I. pseudacorus was reduced from >80% in controls without burial to <40% even at minimal wrack depths of 1 cm. Few I. pseudacorus seedlings were able to emerge from wrack burial of up to 8 cm in depth. We also found greater numbers of both quiescent seeds and germinated seeds that did not emerge from wrack burial. Reduced seedling emergence and increased seed quiescence with wrack burial were primarily explained by a reduction in daily temperature variation within the substrate. No senescent seedlings were observed with any depth of wrack burial. In view of our results, the management of I. pseudacorus invasion will be a long-term challenge, requiring continued control due to persistent seeds in wrack-buried seed banks.

Population Dynamic of the Annual Halophyte Salicornia ramosissima in Salt Pans: Towards a Sustainable Exploitation of Its Wild Populations.

Halophyte species growing under stressful conditions, such as the annual species of the Salicornia genus, have been recognized as a source of metabolites of pharmacological and nutraceutical interest. Therefore, planning the extraction of individual plants from wild populations in a sustainable way is especially important in the case of annual species. We studied the environmental matrix and population dynamic of four Salicornia ramosissima populations growing at two elevations in salt pans under a Mediterranean climate. In elevated areas, S. ramosissima populations presented maximum plant densities of between 628-6288 plants m-2 that remained almost constant until fruiting. In contrast, populations in depressed zones presented five-times greater soil-seed-bank densities and maximum plant densities than populations in elevated zones. In this context, populations in depressed zones lost c. 60% of their maximum plant densities from the end of spring and through summer. In whatever way the environmental matrix seemed to control the population dynamic of S. ramosissima in depressed zones, the effects of a stressful environment would interact with plant densities. In this sense, we recorded the density-dependent mortality for the densest population (max. 51,558 plants m-2). Our results are useful for planning a sustainable harvesting of natural populations of S. ramosissima.

Germination niche breadth of invasive Iris pseudacorus (L.) suggests continued recruitment from seeds with global warming.

PREMISE: Understanding recruitment processes of invasive species is central to conservation and management strategies. Iris pseudacorus, an emergent macrophyte, has established invasive populations across a broad global range, and reduces biodiversity in wetland ecosystems. Climate change is altering germination cues, yet studies on the invasion of wetland macrophytes often ignore germination ecology despite its importance to their establishment and spread. METHODS: We explored germination of seeds from invasive I. pseudacorus populations in California in response to seed coat presence or absence, and several environmental factors. Using experimental results in a thermal time model, we derived germination temperature thresholds. RESULTS: Germination of I. pseudacorus seeds did not require cold or warm stratification, and was not affected by seed coat presence or absence. Germination occurred in the dark, although germinability was two- to threefold times greater under light. At constant temperature, thermal time model estimates included 18.3 ± 1.8°C base germination temperature (Tb$({T}_{b}$ ); 28.2 ± 0.5°C optimal temperature (To$({T}_{o}$ ); and 41.0 ± 1.7°C ceiling temperature (Tc$({T}_{c}$ ). Seeds exposed to 36.0°C achieved over 10% germination, and embryos of ungerminated seeds presented 76% viability. Overall, germinability remained relatively low at constant temperatures (≤25%) but was close to 90% under alternating daily temperatures. CONCLUSIONS: Exposure to diurnally fluctuating temperatures is essential for this species to achieve high germination rates. Our study reveals that I. pseudacorus has a broad germination niche supporting its establishment in a relatively wide range of environments, including at high temperatures more frequent with climate change.

Metal effects on germination and seedling development in closely-related halophyte species inhabiting different elevations along the intertidal gradient.

Seed germination and seedling establishment are very sensitive plant stages to metal pollution. Many halophyte species colonizing salt marshes are able to germinate and establish in highly contaminated habitats and low marsh halophyte species seem to show higher tolerance to metals than high marsh species. We analyzed the effects of copper, zinc and nickel in concentrations up to 2000 µM on seed germination and seedling growth in two closely related species of Sarcocornia, S. perennis, a low marsh species, and S. fruticosa, a high marsh species. Germination of both halophytes was not affected by any metal concentration, and their seedling growth, mainly radicle length, was reduced by increasing metal concentrations. Seedlings of S. perennis showed higher tolerance to the three metals than those of S. fruticosa. Our results are useful for designing ecotoxicological bioassays and planning phytoremediation projects in salt marshes.

Seed bank persistence of a South American cordgrass in invaded northern Atlantic and Pacific Coast estuaries.

Invasive alien plant species impart considerable impacts that contribute to the decline of biodiversity worldwide. The ability of an invasive species to overcome barriers to establish and spread in new environments, and the long-term effects of plant invasions supporting their persistence are keys to invasion success. The capacity of introduced species to form soil seed banks can contribute to their invasiveness, yet few studies of invaders have addressed seed bank dynamics. Improved knowledge of this recruitment process can improve conservation management. We studied temporal and spatial changes in soil seed bank characteristics of the cordgrass Spartina densiflora from two continental invaded ranges. In the Odiel Marshes (Southwest Iberian Peninsula), S. densiflora formed transient seed banks (<1 year). At Humboldt Bay Estuary (California), viable seeds persisted for at least 4 years though the germination percentage fell abruptly after the first year from 29 % to less than 5 % of remaining viable seeds. Total soil seed bank density increased with S. densiflora above-ground cover in both estuaries, pointing to the transient component of the seed bank as a critical component of vegetation dynamics during S. densiflora invasion. Even so, seed densities as high as c. 750 seeds m-2 in Odiel Marshes and c. 12 400 seeds m-2 in Humboldt Bay were recorded in some plots without fruiting S. densiflora plants. S. densiflora spikelet (dispersal unit) density was more than double close to the sediment surface than deeper within soil. Our study shows the importance of evaluating seed banks during the design of invasive species management since seed bank persistence may vary among invaded sites, and can affect the timing and duration required for desired management outcomes.

From physiology to salt marsh management challenges with sea level rise: the case of native Spartina foliosa, invasive S. densiflora and their hybrid.

Sea level rise (SLR) imposes increasing salinity and inundation stresses in salt marshes which simultaneously face invasions by exotic plant species. We aimed to improve and apply knowledge on the ecophysiological responses of halophytes to SLR to conservation management of salt marshes. In a mesocosm experiment, we measured and compared phosphoenolpyruvate carboxylase (PEPC) activity and related functional traits of the California-native Spartina foliosa, invasive S. densiflora and their hybrid S. densiflora × foliosa in response to increasing levels of salinity and inundation. S. foliosa was moderately sensitive to salinity, showing a 57% reduction in PEPC specific activity from freshwater to hypersalinity. This native species compensated for the reduction of PEPC activity with increased salinity through 80% higher enzyme activation by phosphorylation. PEPC functional trait responses of S. foliosa were mostly independent of inundation depth. In view of these results, managers should conserve undeveloped lands for accommodation space above current high tide lines to facilitate colonization of stress-tolerant S. foliosa. Our results on functional responses of PEPC traits recorded high sensitivity to salinity for S. densiflora. This was reflected by 65% lower PEPC specific activity together with increasing accumulation of free proline (+96%) and total proteins (+23%) with elevated salinity. These results suggest prioritized eradication of S. densiflora populations in brackish habitats. Measured PEPC responses support the high stress tolerance of the S. densiflora × foliosa hybrid. PEPC traits for the hybrid were mostly independent of salinity and inundation. The hybrid showed higher PEPC-specific activity than S. foliosa (+70%) and S. densiflora (+15%) in freshwater under intermediate inundation. Results suggest that eradication of the hybrid should be the highest management priority. Our study shows that the responses of key functional physiological traits to environmental stresses serve as biological indicators that can guide ecosystem management practices in a scenario of climate change.

Effects of heavy metal pollution on germination and early seedling growth in native and invasive Spartina cordgrasses.

Seed germination and seedling establishment are the stages most sensitive to abiotic stress in the plant life cycle. We analyzed the effects of copper, zinc and nickel on seed germination and early seedling growth of native Spartina maritima and invasive S. densiflora from polluted and non-polluted estuaries. Germination percentages for either species were not affected by any metal at any tested concentration (up to 2000 µM). However, the increase in metal concentration had negative effects on S. densiflora seedlings. The primary effect was on radicle development, representing initial seedling emergence. Spartina densiflora seedlings emerging from seeds from Tinto Estuary, characterized by high bioavailable metal loads, showed higher tolerance to metals than those from less polluted Odiel and Piedras Estuaries. Comparing our results to metal concentrations in the field, we expect S. densiflora seedling development would be negatively impacted in the most metal-polluted areas in Odiel and Tinto Estuaries.

Interactive effects of salinity and inundation on native Spartina foliosa, invasive S. densiflora and their hybrid from San Francisco Estuary, California.

BACKGROUND AND AIMS: Sea level rise (SLR) associated with climate change is intensifying permanent submersion and salinity in salt marshes. In this scenario, hybridization between native and invasive species may result in hybrids having greater tolerance of abiotic stress factors than their parents. Thus, understanding the responses of native and invasive halophytes and their hybrids to interacting physiological stresses imposed by SLR is key to native species conservation. We analysed how salinity, inundation depth and their interaction impact the functional traits of native and invasive cordgrass species and their hybrid (genus Spartina; Poaceae). METHODS: In a mesocosm experiment, we evaluated interactive stress effects of three inundation depths (4.5, 35.5 and 55 cm) and four aqueous salinities (0.5, 10, 20 and 40 ppt) on 27 functional traits of native Spartina foliosa, invasive S. densiflora and their hybrid S. densiflora × S. foliosa from San Francisco Estuary. KEY RESULTS: The combined effect of salinity and inundation led to synergistic effects on leaf biochemical stress indicators. Spartina foliosa behaved as a stress-tolerant species, with high leaf sodium exudation rate and glycine betaine concentrations that also increased with stress. Spartina foliosa was less sensitive to salinity than S. densiflora and the hybrid but was highly growth-limited in response to increased inundation and salinity. Spartina densiflora was fast-growing in low-stress conditions and tolerated moderate interactive stresses. The hybrid produced more biomass, rhizome reserves and tillers than its parents, even under the most stressful conditions. Transgressivity improved the hybrid's capacity to deal with flooding stress more so than its response to increasing salinity. CONCLUSIONS: Based on our observations, we predict that established populations of both native and invasive cordgrasses will experience reduced vegetative and sexual fitness in response to SLR. In particular, the combined effects of high salinity and deep inundation may decrease floret production in S. densiflora, a key trait for the spread of its invasive populations. In contrast, the hybrid likely will be able to sustain its invasiveness under SLR based on its ability to maintain growth and biomass production under stressful conditions.

Changes to the functional traits of phosphoenolpyruvate carboxylase following hybridization in C-4 halophytes.

Hybridization is a relevant evolutionary mechanism linked to the invasiveness of plant species, but little is known about its effect on enzymatic activities in response to stress. We analyzed the effects of salinity on key mechanistic traits of phosphoenolpyruvate carboxylase (PEPC) enzyme for two hybrid taxa derived from native Spartina maritima (Curtis) Fernald and invasive Spartina densiflora Brongn. in comparison with their parental species. Parental species showed contrasted strategies at the PEPC level to cope with salinity. Spartina maritima showed its physiological optimum at 10 to 40 ppt salinity, with high PEPC activity (per unit leaf soluble protein), in contrast to the lower salinity optimum of 0.5 and 10 ppt for S. densiflora, where highest levels of PEPC apparent specific activity coincided with high light-induced activation of PEPC. Both hybrids showed constant PEPC apparent specific activity from fresh water to hypersalinity and exhibited higher net photosynthesis rates in fresh water than their parents. Spartina maritima × densiflora presented three transgressive PEPC-related traits, being the only taxon able to increase its PEPC activation in darkness at high salinity. Spartina densiflora × maritima showed most PEPC-related traits intermediate between its parents. Inheritance types operating differently in reciprocal hybrids determine key functional traits conditioning their ecological performance.

Transgressivity in Key Functional Traits Rather Than Phenotypic Plasticity Promotes Stress Tolerance in A Hybrid Cordgrass.

Hybridization might promote offspring fitness via a greater tolerance to environmental stressors due to heterosis and higher levels of phenotypic plasticity. Thus, analyzing the phenotypic expression of hybrids provides an opportunity to elucidate further plant responses to environmental stress. In the case of coastal salt marshes, sea level rise subjects hybrids, and their parents, to longer tidal submergence and higher salinity. We analyzed the phenotypic expression patterns in the hybrid Spartina densiflora x foliosa relative to its parental species, native S. foliosa, and invasive S. densiflora, from the San Francisco Estuary when exposed to contrasting salinities and inundations in a mesocosm experiment. 37% of the recorded traits displayed no variability among parents and hybrids, 3% showed an additive inheritance, 37% showed mid-parent heterosis, 18% showed best-parent heterosis, and 5% presented worst-parent heterosis. Transgressivity, rather than phenotypic plasticity, in key functional traits of the hybrid, such as tiller height, conveyed greater stress tolerance to the hybrid when compared to the tolerance of its parents. As parental trait variability increased, phenotypic transgressivity of the hybrid increased and it was more important in response to inundation than salinity. Increases in salinity and inundation associated with sea level rise will amplify the superiority of the hybrid over its parental species. These results provide evidence of transgressive traits as an underlying source of adaptive variation that can facilitate plant invasions. The adaptive evolutionary process of hybridization is thought to support an increased invasiveness of plant species and their rapid evolution.

Seedling Emergence from Seed Banks in Ludwigia hexapetala-Invaded Wetlands: Implications for Restoration.

Soil seed banks play a critical role in the maintenance of wetland plant communities and contribute to revegetation following disturbances. Analysis of the seed bank can therefore inform restoration planning and management. Emergence from seed banks may vary in response to hydrologic conditions and sediment disturbances. To assess the community-level impact of exotic Ludwigia hexapetala on soil seed banks, we compared differences in species composition of standing vegetation among invaded and non-invaded wetlands and the degree of similarity between vegetation and soil seed banks in northern California. To determine potential seed bank recruitment of L. hexapetala and associated plant species, we conducted a seedling emergence assay in response to inundation regime (drawdown vs. flooded) and sediment depth (surface vs. buried). Plant species richness, evenness, and Shannon's H' diversity were substantially lower in standing vegetation at L. hexapetala invaded sites as compared to non-invaded sites. Over 12 months, 69 plant taxa germinated from the seed banks, including L. hexapetala and several other exotic taxa. Seedling density varied among sites, being the highest (10,500 seedlings m-2) in surface sediments from non-invaded sites subjected to drawdown treatments. These results signal the need for invasive plant management strategies to deplete undesirable seed banks for restoration success.

Differential Effects of Increasing Salinity on Germination and Seedling Growth of Native and Exotic Invasive Cordgrasses.

Soil salinity is a key environmental factor influencing germination and seedling establishment in salt marshes. Global warming and sea level rise are changing estuarine salinity, and may modify the colonization ability of halophytes. We evaluated the effects of increasing salinity on germination and seedling growth of native Spartina maritima and invasive S. densiflora from wetlands of the Odiel-Tinto Estuary. Responses were assessed following salinity exposure from fresh water to hypersaline conditions and germination recovery of non-germinated seeds when transferred to fresh water. The germination of both species was inhibited and delayed at high salinities, while pre-exposure to salinity accelerated the speed of germination in recovery assays compared to non-pre-exposed seeds. S. densiflora was more tolerant of salinity at germination than S. maritima. S. densiflora was able to germinate at hypersalinity and its germination percentage decreased at higher salinities compared to S. maritima. In contrast, S. maritima showed higher salinity tolerance in relation to seedling growth. Contrasting results were observed with differences in the tidal elevation of populations. Our results suggest S. maritima is a specialist species with respect to salinity, while S. densiflora is a generalist capable of germination of growth under suboptimal conditions. Invasive S. densiflora has greater capacity than native S. maritima to establish from seed with continued climate change and sea level rise.

Supporting Spartina: Interdisciplinary perspective shows Spartina as a distinct solid genus.

In 2014, a DNA-based phylogenetic study confirming the paraphyly of the grass subtribe Sporobolinae proposed the creation of a large monophyletic genus Sporobolus, including (among others) species previously included in the genera Spartina, Calamovilfa, and Sporobolus. Spartina species have contributed substantially (and continue contributing) to our knowledge in multiple disciplines, including ecology, evolutionary biology, molecular biology, biogeography, experimental ecology, biological invasions, environmental management, restoration ecology, history, economics, and sociology. There is no rationale so compelling to subsume the name Spartina as a subgenus that could rival the striking, global iconic history and use of the name Spartina for over 200 yr. We do not agree with the subjective arguments underlying the proposal to change Spartina to Sporobolus. We understand the importance of both the objective phylogenetic insights and of the subjective formalized nomenclature and hope that by opening this debate we will encourage positive feedback that will strengthen taxonomic decisions with an interdisciplinary perspective. We consider that the strongly distinct, monophyletic clade Spartina should simply and efficiently be treated as the genus Spartina.

Some Like It Hot: Maternal-Switching With Climate Change Modifies Formation of Invasive Spartina Hybrids.

Climate change can induce temporary, spatial or behavioral changes in species, so that only some species can adapt to the new climatic conditions. In the case of invasive species, it is expected that they will be promoted in a context of global change, given their high tolerance to environmental factors and phenotypic plasticity. Once in the invaded range, these species can hybridize with native species thus introducing their genotype in the native biota. However, the effects that climate change will have on this process of invasion by hybridization remain unclear. We evaluated the historical establishment of the reciprocal hybrids between the native Spartina maritima and the invasive S. densiflora in the Gulf of Cadiz (SW Iberian Peninsula) and we related it to climatic changes during the period 1955-2017. Our results showed that, according to their dating based on their rate of lateral expansion rates, the establishment of S. maritima × densiflora and S. densiflora × maritima in the Gulf of Cadiz has occurred in the last two centuries and has been related to changes in air temperature and rainfall during the flowering periods of their parental species, with antagonist impacts on both hybrids. Thus, the hybrid S. densiflora × maritima has been established in years with mild ends of spring and beginning of summer when the flowering of S. maritima lengthened and its pollen production was higher, and it coincided with the beginning of the flowering period of S. densiflora. Moreover, the establishment of this hybrid was related to higher spring/summer rainfalls, probably due to the reduction in salinity in middle marshes. However, the hybrid S. maritima × densiflora, was established mainly in warmer spring/summers in which the proportion of pollen:ovule of S. maritima was reduced favoring its pollination by S. densiflora. As a consequence of the promotion of S. maritima × densiflora with climate change, the native and endangered species S. maritima would be threatened, as both taxa share the same habitat and the hybrid shows a remarkably higher competitive potential.

Phenotypic plasticity of polyploid plant species promotes transgressive behaviour in their hybrids.

Hybridization is a frequent process that leads to relevant evolutionary consequences, but there is a lack of studies regarding the relationships of the variability of the response of parental plant species to environmental gradients and the responses of their hybrids at a phenotypic level. We designed an experiment in which we exposed two reciprocal cordgrass hybrids, Spartina maritima × densiflora and S. densiflora × maritima, and their parental species to four salinity concentrations for 30 days. The main objectives were to compare the performance of the hybrids with that of their parents, to distinguish the phenotypic inheritance operating in the hybrids and to analyse the relationships between the variability in the responses of the parents and the responses of their hybrids to salinity. We characterized the responses and the degree of variability for 37 foliar traits. Both hybrids presented greater salinity tolerance than their parents, showing their highest percentage of transgressive traits at both extremes of the salinity gradient. When the parental plants themselves showed a more plastic response for a given trait, there was a greater chance that their hybrid developed a transgressive behaviour for this trait. This finding supports a new focus to be applied for the artificial development of vigorous hybrid crops.

Low genetic diversity contrasts with high phenotypic variability in heptaploid Spartina densiflora populations invading the Pacific coast of North America.

Species can respond to environmental pressures through genetic and epigenetic changes and through phenotypic plasticity, but few studies have evaluated the relationships between genetic differentiation and phenotypic plasticity of plant species along changing environmental conditions throughout wide latitudinal ranges. We studied inter- and intrapopulation genetic diversity (using simple sequence repeats and chloroplast DNA sequencing) and inter- and intrapopulation phenotypic variability of 33 plant traits (using field and common-garden measurements) for five populations of the invasive cordgrass Spartina densiflora Brongn. along the Pacific coast of North America from San Francisco Bay to Vancouver Island. Studied populations showed very low genetic diversity, high levels of phenotypic variability when growing in contrasted environments and high intrapopulation phenotypic variability for many plant traits. This intrapopulation phenotypic variability was especially high, irrespective of environmental conditions, for those traits showing also high phenotypic plasticity. Within-population variation represented 84% of the total genetic variation coinciding with certain individual plants keeping consistent responses for three plant traits (chlorophyll b and carotenoid contents, and dead shoot biomass) in the field and in common-garden conditions. These populations have most likely undergone genetic bottleneck since their introduction from South America; multiple introductions are unknown but possible as the population from Vancouver Island was the most recent and one of the most genetically diverse. S. densiflora appears as a species that would not be very affected itself by climate change and sea-level rise as it can disperse, establish, and acclimate to contrasted environments along wide latitudinal ranges.

Competition from native hydrophytes reduces establishment and growth of invasive dense-flowered cordgrass (Spartina densiflora).

Experimental studies to determine the nature of ecological interactions between invasive and native species are necessary for conserving and restoring native species in impacted habitats. Theory predicts that species boundaries along environmental gradients are determined by physical factors in stressful environments and by competitive ability in benign environments, but little is known about the mechanisms by which hydrophytes exclude halophytes and the life history stage at which these mechanisms are able to operate. The ongoing invasion of the South American Spartina densiflora in European marshes is causing concern about potential impacts to native plants along the marsh salinity gradient, offering an opportunity to evaluate the mechanisms by which native hydrophytes may limit, or even prevent, the expansion of invasive halophytes. Our study compared S. densiflora seedling establishment with and without competition with Phragmites australis and Typha domingensis, two hydrophytes differing in clonal architecture. We hypothesized that seedlings of the stress tolerant S. densiflora would be out-competed by stands of P. australis and T. domingensis. Growth, survivorship, biomass patterns and foliar nutrient content were recorded in a common garden experiment to determine the effect of mature P. australis and T. domingensis on the growth and colonization of S. densiflora under fresh water conditions where invasion events are likely to occur. Mature P. australis stands prevented establishment of S. densiflora seedlings and T. domingensis reduced S. densiflora establishment by 38%. Seedlings grown with P. australis produced fewer than five short shoots and all plants died after ca. 2 yrs. Our results showed that direct competition, most likely for subterranean resources, was responsible for decreased growth rate and survivorship of S. densiflora. The presence of healthy stands of P. australis, and to some extent T. domingensis, along river channels and in brackish marshes may prevent the invasion of S. densiflora by stopping the establishment of its seedlings.

Phenotypic plasticity of invasive Spartina densiflora (Poaceae) along a broad latitudinal gradient on the Pacific Coast of North America.

PREMISE OF THE STUDY: Phenotypic acclimation of individual plants and genetic differentiation by natural selection within invasive populations are two potential mechanisms that may confer fitness advantages and allow plants to cope with environmental variation. The invasion of Spartina densiflora across a wide latitudinal gradient from California (USA) to British Columbia (Canada) provides a natural model system to study the potential mechanisms underlying the response of invasive populations to substantial variation in climate and other environmental variables. METHODS: We examined morphological and physiological leaf traits of Spartina densiflora plants in populations from invaded estuarine sites across broad latitudinal and climate gradients along the Pacific west coast of North America and in favorable conditions in a common garden experiment. KEY RESULTS: Our results show that key foliar traits varied widely among populations. Most foliar traits measured in the field were lower than would be expected under ideal growing conditions. Photosynthetic pigment concentrations at higher latitudes were lower than those observed at lower latitudes. Greater leaf rolling, reduced leaf lengths, and lower chlorophyll and higher carbon concentrations were observed with anoxic sediments. Lower chlorophyll to carotenoids ratios and reduced nitrogen concentrations were correlated with sediment salinity. Our results suggest that the variations of foliar traits recorded in the field are a plastic phenotypic response that was not sustained under common garden conditions. CONCLUSIONS: SPARTINA DENSIFLORA shows wide differences in its foliar traits in response to environmental heterogeneity in salt marshes, which appears to be the result of phenotypic plasticity rather than genetic differentiation.

Native plant restoration combats environmental change: development of carbon and nitrogen sequestration capacity using small cordgrass in European salt marshes.

Restoration of salt marshes is critical in the context of climate change and eutrophication of coastal waters because their vegetation and sediments may act as carbon and nitrogen sinks. Our primary objectives were to quantify carbon (C) and nitrogen (N) stocks and sequestration rates in restored marshes dominated by Spartina maritima to provide support for restoration and management strategies that may offset negative aspects of eutrophication and climate change in estuarine ecosystems. Sediment C content was between ca. 13 mg C g(-1)and sediment N content was ca. 1.8 mg N g(-1). The highest C content for S. maritima was recorded in leaves and stems (ca. 420 mg C g(-1)) and the lowest in roots (361 ± 4 mg C g(-1)). S. maritima also concentrated more N in its leaves (31 ± 1 mg N g(-1)) than in other organs. C stock in the restored marshes was 29.6 t C ha(-1); ca. 16 % was stored in S. maritima tissues. N stock was 3.6 t N ha(-1), with 8.3 % stored in S. maritima. Our results showed that the S. maritima restored marshes, 2.5 years after planting, were sequestering atmospheric C and, therefore, provide some mitigation for global warming. Stands are also capturing nitrogen and reducing eutrophication. The concentrations of C and N contents in sediments, and cordgrass relative cover of 62 %, and low below-ground biomass (BGB) suggest restored marshes can sequester more C and N. S. maritima plantations in low marshes replace bare sediments and invasive populations of exotic Spartina densiflora and increase the C and N sequestration capacity of the marsh by increasing biomass production and accumulation.