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.

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.

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.

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.