Soil physicochemical properties associated with the yield and phytochemical composition of the edible halophyte Crithmum maritimum.

There is growing interest in the consumption of halophytes due to their excellent nutritional profile and antioxidant properties, and their cultivation offers viable alternatives in the face of irreversible global salinization of soils. Nevertheless, abiotic factors strongly influence their phytochemical composition, and little is known about how growing conditions can produce plants with the best nutritional and functional properties. Crithmum maritimum is an edible halophyte with antioxidant properties and considerable potential for sustainable agriculture in marginal environments. However, it is found naturally in contrasting habitats with variable soil physicochemical properties and the extent to which edaphic factors can influence plant performance, accumulation of phytochemicals and their quality remains unknown. We investigated the influence of soil physicochemical properties (texture, pH, electrical conductivity, organic matter content and mineral element concentrations) on growth and reproductive performance, nutritional traits, and the accumulation of specific metabolites in C. maritimum. Soil, leaf and seed samples were taken from eight C. maritimum populations located on the southern coasts of Spain and Portugal. We found greater vegetative growth and seed production in coarser, sandier soils with lower microelement concentrations. The nutritional traits of leaves varied, with soil organic matter and macronutrient content associated with reduced leaf Na, protein and phenolic (mainly flavonoid) concentrations, whereas soils with lower pH and Fe concentrations, and higher clay content yielded plants with lower leaf Zn concentration and greater accumulation of hydroxycinnamic acids. The nutritional value of the seed oil composition appeared to be enhanced in soils with coarser texture and lower microelement concentrations. The accumulation of specific phenolic compounds in the seed was influenced by a wide range of soil properties including texture, pH and some microelements. These findings will inform the commercial cultivation of C. maritimum, particularly in the economic exploitation of poorly utilized, saline soils.

Disentangling elevation, annual flooding regime and salinity as hydrochemical determinants of halophyte distribution in non-tidal saltmarsh.

BACKGROUND AND AIMS: Hydrological disconnection, especially in a Mediterranean climate, creates coastal saltmarshes with an annual cycle of flooding that are unlike tidally inundated systems. Winter rainfall produces long, continuous hydroperiods, alternating with continuous exposure caused by evaporation in warm, rain-free summers. We aimed to distinguish the effects of elevation, hydroperiod and salinity on annual and perennial halophytes in such a system. METHODS: We recorded vegetation and sediment salinity in permanent quadrats on a marsh in the Doñana National Park, Spain, over seven consecutive years with widely differing rainfall. Elevation was determined from LIDAR data and the duration of the annual hydroperiod from satellite imagery. The independent effects of collaterally varying elevation, hydroperiod and salinity on species distribution were examined using generalized linear models and hierarchical partitioning. KEY RESULTS: Both hydroperiod and salinity were inversely related to elevation but interannual fluctuations in rainfall facilitated discrimination of independent effects of the three collaterally varying factors on halophyte distribution. Perennial distribution was strongly structured by elevation, whereas many annual species were more sensitive to hydroperiod. The independent effects of salinity varied according to individual species' salt tolerance from positive to negative. Thus life-history and, in the case of annuals, phenology were important in determining the relative impact of elevation and hydroperiod. CONCLUSIONS: The consequences of elevation for halophyte distribution in seasonally flooded saltmarshes are fundamentally different from those in tidal marshes, because protracted and frequent flooding regimes require different adaptations, and because of the unpredictability of flooding from year to year. These differences could explain greater species diversity in non-tidal marshes and the absence of key saltmarsh species prominent in tidal marshes. The vegetation of non-tidal marshes will be particularly susceptible to the more extreme annual cycles of temperature and rainfall predicted for Mediterranean climates.

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.

Is saltmarsh restoration success constrained by matching natural environments or altered succession? A test using niche models.

Restored habitats, such as saltmarsh created through managed realignment, sometimes fail to meet targets for biological equivalence with natural reference sites. Understanding why this happens is important in order to improve restoration outcomes.Elevation in the tidal frame and sediment redox potential are major controls on the distribution of saltmarsh plants. We use niche models to characterize 10 species' responses to these, and test whether differences in species occurrence between restored and natural saltmarshes in the UK result from failure to recreate adequate environmental conditions.Six species occurred less frequently in recently restored marshes than natural marshes. Failure of restored marshes to achieve the elevation and redox conditions of natural marshes partially explained the underrepresentation of five of these species, but did not explain patterns of occurrence on older (>50 years) restored marshes.For all species, an effect of marsh age remained after controlling for differences in environmental conditions. This could be due to differences in successional mechanism between restored and natural marshes. In recently restored marshes, high-marsh species occurred lower in the tidal frame and low-marsh species occurred higher in the tidal frame than in natural marshes. This supports the hypothesis that competition is initially weaker in restored marshes, because of the availability of bare sediment across the whole tidal frame. Species that establish outside their normal realized niche, such as Atriplex portulacoides, may inhibit subsequent colonization of other species that occurred less frequently than expected on older restored marshes. Synthesis and applications. Niche models can be used to test whether abiotic differences between restored sites and their natural counterparts are responsible for discrepancies in species occurrence. In saltmarshes, simply replicating environmental conditions will not result in equivalent species occurrence.

A re-examination of the Salicornias (Amaranthaceae) of Saudi Arabia and their polymorphs.

During the period from 1964 to 1999 Saudi Arabian species of Salicornia were wrongly treated under the European species, S. europaea L. Recent explorations proved that there are two separate allopatric species of Salicornia in Saudi Arabia, one inhabiting the inland salt-marshes of the Najd (highlands) and the other inhabiting the Arabian Gulf Coast (lowlands). Morphological, ecological and exploratory studies confirm that they are two distinct species. The two species differ in features of bark, axillary spikes, basal vegetative segment(s) of spike, fertile segments, colour of senescent plants, and flowering, fruiting and germination phenology. As both the species have been described earlier from Iran, they are now new records for Saudi Arabia. The species are, S. persica ssp. iranica (Akhani) Kadereit & Piirainen and S. sinus-persica Akhani. S. sinus-persica, of which the status was thought doubtful has been confirmed. Both the species have been described and illustrated. Each species comprises a number of polymorphs. As leaves and flowers are rudimentary, confusing species circumscriptions, a proliferation of binomials has resulted in the taxonomy of Salicornia. To mitigate such confusion, the full range of variability of the Saudi Arabian species has been documented.

Potential disruption of pollination in a sexually deceptive orchid by climatic change.

Warmer springs advance many phenological events, including flowering time in plants and the flight time of insects. Pollination by insects, an ecosystem service of immense economic and conservation importance, depends on synchrony between insect activity and flowering time. If plants and their pollinators show different phenological responses to climate warming, pollination could fail. Information about the effects of warming on specific plant-insect mutualisms is difficult to obtain from complex pollination networks. In contrast, the extraordinarily specific deceptions evolved by orchids that attract a very narrow range of pollinators allow direct examination of the potential for climatic warming to disrupt synchrony. Here we show that a sexually deceptive orchid and the solitary bee on which it depends for pollination will diverge in phenology with increasing spring temperature. Male bees inadvertently pollinate the orchid flowers during pseudocopulation. Analysis of museum specimens (1893-2007) and recent field-based records (1975-2009) showed that flight date of the solitary bee Andrena nigroaenea is advanced more by higher temperatures than is flowering date in the deceptive orchid Ophrys sphegodes. Male bees emerged slightly earlier than females, which attract male copulatory attentions away from the deceptive flowers. Warming by as little as 2°C increased both the probability of male flight and the proportion of females flying in the bee population before orchid flowering; this would reduce the frequency of pseudocopulation and thus lower pollination success rate in the orchid. Our results demonstrate a significant potential for coevolved plant-pollinator relationships to be disrupted by climatic warming.

Silicon alleviates deleterious effects of high salinity on the halophytic grass Spartina densiflora.

The non-essential element silicon is known to improve plant fitness by alleviating the effects of biotic and abiotic stresses, particularly in crops. However, its possible role in the exceptional tolerance of halophytes to salinity has not been investigated. This study reports the effect of Si supply on the salinity tolerance of the halophytic grass Spartina densiflora; plants were treated with NaCl (0-680 mM), with or without silicon addition of 500 µM, in a glasshouse experiment. Plant responses were examined using growth analysis, combined with measurements of gas exchange, chlorophyll fluorescence and photosynthetic pigment concentrations. In addition, tissue concentrations of aluminium, calcium, copper, iron, potassium, magnesium, sodium, phosphorus and silicon were determined. Although high salinity decreased growth, this effect was alleviated by treatment with Si. Improved growth was associated with higher net photosynthetic rate (A), and greater water-use efficiency (WUE). Enhanced A at high salinity could be explained by beneficial effects of Si on the photochemical apparatus, and on chlorophyll concentrations. Ameliorative effects of Si were correlated with reduced sodium uptake, which was unrelated to a reduction in the transpiration rate, since Si-supplemented plants had higher stomatal conductances (G(s)). These plants also had higher tissue concentrations of essential nutrients, suggesting that Si had a positive effect on the mineral nutrient balance in salt-stressed plants. Si appears to play a significant role in salinity tolerance even in a halophyte, which has other, specific salt-tolerance mechanisms, through diverse protective effects on the photosynthetic apparatus, water-use efficiency and mineral nutrient balance.

Water quality as a threat to aquatic plants: discriminating between the effects of nitrate, phosphate, boron and heavy metals on charophytes.

• Eutrophication is a threat to wetlands worldwide. Elevated phosphorus concentration is often the main driver of loss of biodiversity and ecosystem function, but effects of phosphorus and nitrogen have proved largely inseparable, because they vary colinearly. Charophytes, aquatic algae that are very close to the evolutionary link with the land-plant lineage, provide a sensitive system for disentangling complex pollutant threats. • Here, we investigated aquatic vegetation and water quality at the principal sites for charophyte biodiversity in the UK and used hierarchical partitioning to discriminate independent effects of pollutants on their occurrence. A laboratory experiment examined the growth responses of a representative species (Chara globularis) to nitrate. • Nitrate-N exerted the greatest detrimental effect on charophyte occurrence in the field. Furthermore, growth of C. globularis in the laboratory was inhibited above very low concentrations. Smaller independent effects of copper (Cu), cadmium (Cd), cobalt (Co), phosphate-P, nickel (Ni), boron (B) and manganese (Mn) on charophyte occurrence were discriminated. • It is possible to separate the deleterious effects of phosphorus and nitrogen on aquatic organisms in the field. Nitrate is a critical factor and a mean annual average concentration limit of c. 2 mg l⁻¹ nitrate-N is necessary to protect charophytes and their services within wetland ecosystems.

Carry-over of differential salt tolerance in plants grown from dimorphic seeds of Suaeda splendens.

BACKGROUND AND AIMS: Halophytic species often show seed dimorphism, where seed morphs produced by a single individual may differ in germination characteristics. Particular morphs are adapted to different windows of opportunity for germination in the seasonally fluctuating and heterogeneous salt-marsh environment. The possibility that plants derived from the two morphs may also differ physiologically has not been investigated previously. METHODS: Experiments were designed to investigate the germination characteristics of black and brown seed morphs of Suaeda splendens, an annual, C(4) shrub of non-tidal, saline steppes. The resulting seedlings were transferred to hydroponic culture to investigate their growth and photosynthetic (PSII photochemistry and gas exchange) responses to salinity. KEY RESULTS: Black seeds germinated at low salinity but were particularly sensitive to increasing salt concentrations, and strongly inhibited by light. Brown seeds were unaffected by light, able to germinate at higher salinities and generally germinated more rapidly. Ungerminated black seeds maintained viability for longer than brown ones, particularly at high salinity. Seedlings derived from both seed morphs grew well at high salinity (400 mol m(-3) NaCl). However, seedlings derived from brown seeds performed poorly at low salinity, as reflected in relative growth rate, numbers of branches produced, F(v)/F(m) and net rate of CO(2) assimilation. CONCLUSIONS: The seeds most likely to germinate at high salinity in the Mediterranean summer (brown ones) retain a requirement for higher salinity as seedlings that might be of adaptive value. On the other hand, black seeds, which are likely to delay germination until lower salinity prevails, produce seedlings that are less sensitive to salinity. It is not clear why performance at low salinity, later in the life cycle, might have been sacrificed by the brown seeds, to achieve higher fitness at the germination stage under high salinity. Analyses of adaptive syndromes associated with seed dimorphism may need to take account of differences over the entire life cycle, rather than just at the germination stage.

Growth and photosynthetic responses to salinity of the salt-marsh shrub Atriplex portulacoides.

BACKGROUND AND AIMS: Atriplex (Halimione) portulacoides is a halophytic, C(3) shrub. It is virtually confined to coastal salt marshes, where it often dominates the vegetation. The aim of this study was to investigate its growth responses to salinity and the extent to which these could be explained by photosynthetic physiology. METHODS: The responses of young plants to salinity in the range 0-700 mol m(-3) NaCl were investigated in a glasshouse experiment. The performance of plants was examined using classical growth analysis, measurements of gas exchange (infrared gas analysis), determination of chlorophyll fluorescence characteristics (modulated fluorimeter) and photosynthetic pigment concentrations; total ash, sodium, potassium and nitrogen concentrations, and relative water content were also determined. KEY RESULTS: Plants accumulated Na(+) approximately in proportion to external salinity. Salt stimulated growth up to an external concentration of 200 mol m(-3) NaCl and some growth was maintained at higher salinities. The main determinant of growth response to salinity was unit leaf rate. This was itself reflected in rates of CO(2) assimilation, which were not affected by 200 mol m(-3) but were reduced at higher salinities. Reductions in net photosynthetic rate could be accounted for largely by lower stomatal conductance and intercellular CO(2) concentration. Apart from possible effects of osmotic shock at the beginning of the experiment, salinity did not have any adverse effect on photosystem II (PSII). Neither the quantum efficiency of PSII (Phi(PSII)) nor the chlorophyll fluorescence ratio (F(v)/F(m)) were reduced by salinity, and lower mid-day values recovered by dawn. Mid-day F(v)/F(m) was in fact depressed more at low external sodium concentration, by the end of the experiment. CONCLUSIONS: The growth responses of the hygro-halophyte A. portulacoides to salinity appear largely to depend on changes in its rate of photosynthetic gas exchange. Photosynthesis appears to be limited mainly through stomatal conductance and hence intercellular CO(2) concentration, rather than by effects on PSII; moderate salinity might stimulate carboxylation capacity. This is in contrast to more extreme halophytes, for which an ability to maintain leaf area can partially offset declining rates of carbon assimilation at high salinity.

Significance of sighting rate in inferring extinction and threat.

We are now entering a time of immense environmental upheaval in which, increasingly, experts are required to provide conservation assessments. Quantitative assessment of trends in species' range and abundance is costly, requiring extensive field studies over a long period of time. Unfortunately, many species are only known through a few "chance" sightings or a handful of specimens, and therefore extinction may be even harder to ascertain. Several methods have been proposed for estimating the probability of extinction. However comparison within and between species is difficult because of variations in sighting rates. We applied a probabilistic method that incorporates sighting rate to the sighting record of Vietnamese slipper orchids (Paphiopedilum). The method generates a probability that another sighting will occur given the previous sighting rate and the time since last observation. This allows greater comparability between species discovered at different times. Its predictions were more highly correlated with the World Conservation Union criteria than previous methods. Trends in data collection and the political climate of a country, which affects access to material, are important potential sources of variation that affect sighting rates. A lack of understanding of the process by which data are generated makes inferring extinction from sighting records difficult because extinction status depends on how the sighting rate varies. However such methods allow rapid conservation prioritization of taxa that are poorly known and would otherwise go unassessed.