Patterns of variation in morphological characters and isoenzymes in populations of Sphagnum capillifolium (Ehrh.) Hedw. and S. rubellum Wils. from two bogs in southern Sweden

Abstract

Isoenzymatic and morphological data of four populations from two bogs in southern Sweden are presented. From each bog one population of Sphagnum rubellum was sampled from the mire expanse and one population of S. capillifolium from the mire margin. Data from isoenzyme-electrophoresis show that there is agenetic difference corresponding to the two morphological taxa. It is also indicated that they may hybridize. The fact that the taxa seem to be better separated at the continental Aggarps Mosse than at the suboceanic Traneröds Mosse would suggest that hybridization occurs more often in coastal than in inland climates.     

Population structure and interspecific differentiation of the peat moss sister speciesSphagnum rubellum andS. capillifolium (Sphagnaceae) in northern Europe

Isozyme electrophoresis was used to study the morphologically similar sister speciesSphagnum rubellum andS. capillifolium from a sample of 1313 plants representing 37 populations from Scandinavia, Great Britain and S Germany. The mean pairwise genetic identities (I) among conspecific populations were 0.976 forS. rubellum and 0.969 forS. capillifolium, versus 0.627 between populations of the two species. Interspecific gene flow was indicated by the observation of occasional plants in sympatric populations with alleles otherwise unique to the other species. Populations of bisexualS. capillifolium were significantly more variable than populations of unisexualS. rubellum. Alpine populations ofS. rubellum andS. capillifolium were dominated by few genotypes, and differentiation among populations was pronounced, indicating a low level of sexual recombination. InS. rubellum, maximum variability was found in western areas with high annual precipitation. Distribution of alleles inS. rubellum indicated restricted gene flow between Great Britain and Scandinavia. Postglacial migration from separate refugia may explain large-scale variation inS. rubellum.     

Genetic diversity in populations of Sphagnum capillifolium from the mountains of Bulgaria,and their possible refugial role

Abstract

Genetic diversity in eight populations of Sphagnum capillifolium from different Bulgarian mountains was investigated by means of isozyme electrophoresis. High levels of allelic diversity were found (H_S = 0.119), comparable to earlier estimates for northern European populations (H_S = 0.116). Strong differentiation among populations and a low number of widespread genotypes suggest a high degree of isolation and restricted gene flow between populations, which is consistent with generally small and scattered populations. The large proportion of distinguishable genotypes (mean 0.498) suggests high levels of out-crossing either currently or in the past. Introgression between S. capillifolium and S. rubellum, a species not found in Bulgaria, was suggested by the occurrence of rubellum-alleles in five populations from different mountains. This could be explained by an ancient hybridization event in a sympatric population. Based on (1) the high genetic diversity, (2) the fairly wide distribution of alien alleles, and (3) the isolated distribution of populations even within one mountain, a possible survival of S. capillifolium in the Balkan area during the Quaternary ice periods is hypothesized.     

Sphagnum subfulvum Sjörs in Ireland compared with the occurrences in Norway

Abstract

The discovery of Sphagnum subfulvum in Western Ireland is reported and the Irish specimens are described. The habitat of the species at Errisbeg, Co. Galway, is described and compared with the occurrences in Norway. It is concluded that the ecology of S. subfulvum at Errisbeg closely resembles its ecology in Norway where it is an intermediate/rich fen species, and in the ‘mud-bottomto hummock’ vegetational gradient it belongs to lawn and low hummock vegetation. The occurrence of Sphagnum subfulvum on flushed mountain slopesin West Mayo is apparently related to localised basic conditions as indicated by its association with Schoenus nigricans, Anagallis tenella, Selaginella selaginoides and Breutelia chrysocoma. Sphagnum subfulvum is reported from raised bog in Co. Roscommon. Its occurrence in ombrotrophic vegetation is somewhat unexpected but the ombrotrophic vegetation in Western Ireland includes some species which are restricted to minerotrophic vegetation elsewhere.     

Taxon delimitation and genetic similarities of the Sphagnum imbricatum complex,as revealed by enzyme electrophoresis

Abstract

Isozyme electrophoretic studies were used to assess genetic variation in the Sphagnum imbricatum complex in a sample of 1332 plants from 39 populations from western Europe, eastern North America and Japan. Mean pairwise genetic distance among populations clustered and depicted in a UPGMA phenogram correspond to the recognition of four species, viz. Sphagnum portoricense , S. affine , S. imbricatum and S. austinii. The mean pairwise genetic identity among conspecific populations were 0.976, 0.847 and 0.841 for S. austinii , S. affine and S. portoricense, respectively. The mean pairwise genetic identity among taxa was 0.525 (S. austinii–S. affine), 0.476 (S. affine –S. portoricense), 0.600 (S. affine–S. imbricatum), 0.484 (S. imbricatum–S. austinii), 0.629 (S. imbricatum –S. portoricense) and 0.285 (S. austinii–S. portoricense). Populations of S. austinii in Europe are found to be genetically eroded (Hs = 0.001 ± 0.000), (P₉₅ = 0.00), probably due to severe bottlenecks caused by a series of founder effects during postglacial migration from a limited number of glacial refugia in S.W. Europe. The mean genetic diversity of S. affine (Hs = 0.122 ± 0.020) is at the same level as up to now reported for the more variable congeneric species. Among individuals of S. affine, 0.27% displayed mixed markers, indicating that, on rare occasions, hybridization may occur between S. affine and S. austinii. Preliminary genetic analysis of S. steerei supports the recognition of this taxon.     

Habitat-specific genetic effects on growth rate and morphology across pH and water-level gradients within a population of the moss Sphagnum angustifolium (Sphagnaceae)

To study genetic adaptations in bryophytes on small ecological and spatial scales and to assess the adaptive significance of morphological trait variation, genotypes of Sphagnum angustifolium originating from habitats characterized by different pH and height above water table were clonally propagated and grown along the same gradients that exist in the field. Clones from ombrotrophic habitats grew consistently better ombrotrophically than clones from minerotrophic habitats and vice versa, suggesting that the genotypes were adapted to different pH levels. Genetic variation was found in several morphological traits, but habitat-specific genetic effects were detected only in length of spreading branches. Covariation between morphology and growth was generally environmentally induced. Positive and negative cross-environment genetic correlations suggested the presence of constraints on adaptive reaction norm evolution. The indications of small-scale genetic adaptations suggest either selective establishment of genotypes adapted to specific habitats, strong selective forces operating at the later stages of the life cycle, restricted gene flow over short distances, or a combination of these. In contrast to prevailing views, these results indicate that bryophytes are likely to respond genetically to small-scale environmental gradients.     

Sphagnum divinum (sp. nov.) and S. medium Limpr. and their relationship to S. magellanicum Brid.

Sphagnum magellanicum has been viewed as being a predominantly circumpolar species in the northern hemisphere, but it occurs in the southern hemisphere and was originally described from the southern parts of Chile. It is an ecologically important species in mire ecosystems and has been extensively used as a model to study processes of growth, carbon sequestration and peat decomposition. Molecular and experimental studies have, however, revealed genetic structure within S. magellanicum, and morphological differences associated with these genetic groups. Here we describe Sphagnum divinum in Sphagnum subgenus Sphagnum (Sphagnaceae, Bryophyta) as a new species, based on molecular and morphological evidence. Sphagnum medium is reinstated as a distinct species and is epitypified. Consequently, a new species concept of S. magellanicum is presented including an epitypification. Important morphological characters to separate these three species in the field and under the microscope are presented. Ecology and distribution differ among the species; S. divinium has a wide habitat range including mire margin, forested peatlands and moist heaths, and a circumpolar distribution around the northern hemisphere. Sphagnum medium seems to be more restricted to ombrotrophic mire expanse habitats and shows an amphi-Atlantic distribution in the northern hemisphere. Sphagnum magellanicum has a very broad ecological niche in peatlands and is found in most mire habitats in Tierra del Fuego on the southern tip of South America.     

Expansion of Sphagnum fallax in bogs: striking the balance between N and P availability

Abstract

Nitrogen deposition may cause shifts in the Sphagnum species composition of bogs, ultimately affecting the conservation value of these systems. We studied the effects of N and P on the expansion of S. fallax and S. flexuosum in bogs. We related historical census data of S. fallax, S. flexuosum, and four of their accompanying species to changes in N deposition. In addition, we conducted two fertilization experiments with N and P; one at a low deposition site with S. flexuosum and one at two high deposition sites with S. fallax. Finally, we related existing data on capitulum N and P concentrations of S. fallax to its abundance in the field.

A relative increase in observed frequency of S. fallax coincided with an historical increase in N deposition in the Netherlands. There was no indication that S. fallax consistently outcompeted one of the other five Sphagnum species; the observed frequency of the Sphagnum species analysed was rather stable with time. The census data on S. flexuosum did not indicate a response to N deposition, but the species expanded at the low N deposition site when extra N was applied. In contrast, the expansion of S. fallax at the high deposition sites was limited by P. Organic nutrient concentrations suggested that when S. fallax can maintain a capitulum N concentration of 7 mg g^-1 or higher and a P concentration of 0.7 mg g^-1 or higher the species can grow to dominate.

We conclude that S. fallax will gradually colonize an increasing number of new habitats in areas with a low, albeit increasing, N deposition, but will only grow to dominate when P supply is adequate. Then, the expansion of S. fallax may lead to ousting of the other Sphagnum species present.     

Phenotypic variation in the Sphagnum recurvum complex: a cultivation experiment

Abstract

Three morphs in the Sphagnum recurvum complex sampled from two mire localities in Central Norway were cultivated under different water levels and nutrient conditions in order to assess the amount and direction of phenotypic plasticity in several morphological variables. The factor ‘morph’ accounted for much of the variation. However, many of the shoot, branch, leaf and cell variables responded to the experimentally altered conditions, and water level was particularly important. Only the spreading:hanging branch length ratio and various stem leaf variables distinguished the Sphagnum angustifolium moiph from the S. fallax and S. isoviitae morphs regardless of environmental conditions and sampling locality. The results indicated that there is a continuum of genotypes shared by S. fallax and S. isoviitae.     

Effects of water chemistry,nutrient supply and interspecific interactions on the replacement of Sphagnum subnitens by S. fallax in fens

Abstract

In Dutch rich fens, mid-successional stages dominated by Sphagnum subnitens are rapidly being replaced by other vegetation, dominated by S. fallax.

In both species growth was strongly reduced in more mineral-rich ground water and simulated river water, compared to growth in rain water.

Both species were positively affected by added phosphorus supply, whereas added supply of nitrogen did not alter growth rates.

Nutrient supply seemed to compensate for the intolerance to high mineral levels in S. fallax: nutrient addition stimulated growth in length in ground water and simulated river water. Such compensation was not found in S. subnitens.

Growth in mixed culture positively affected Sphagnum subnitens and negatively affected its successor S. fallax.

The success of Sphagnum fallax in early stages of succession in more eutrophic fens may to some extent be explained by its tolerance to high mineral levels with increased nutrient supply.     

Two new African species of Lejeuneaceae

Abstract

At present, the available information about the liverworts of Sphagnum magellanicum Brid. mires from South America is very limited. The main aim of this paper is to present a comprehensive study of the liverwort flora of this habitat, as well as details on the chorology and ecology of the taxa involved. Four different environmental units were examined, based on the water table level and plant cover, in three representative raised bogs from Tierra del Fuego National Park Ushuaia. Floristic relevs were recorded in 93 square plots 050050m², distributed at random in the environmental units. Twentyfour liverwort species have been identified the majority have a Subantarctic distribution and there were a remarkable number of Magellanean endemics 11 species. Among the rare species, there were two new records for the Argentinian Province of Tierra del Fuego, Cephaloziella byssacea and Chiloscyphus notophyllus. The ecological preferences of the species were analysed based on their frequencies in the four environmental units examined. Only six species demonstrated significant correlation with some of the environmental units. Riccardia pallidevirens, R. georgiensis, R. alcicornis, and Cephaloziella varians, showed a preference for waterlogged and wet habitats, while Calypogeia sphagnicola and Cephalozia chilensis were linked to dense carpets of Sphagnum magellanicum.     

A New Approach for Tracking Vegetation Change after Restoration: A Case Study with Peatlands

Developing objective tools for tracking progress of restored sites is of general concern. Here, we present an innovative approach based on principal response curves (PRC) and species classification according to their preferential habitats to monitor changes in community composition. Following large‐scale restoration of a cut‐over peatland, vegetation was surveyed biannually over 8 years. We evaluated whether the establishing plant communities fell within the range of natural variation. We used both general diversity curves and PRC applied on plant species grouped by preferred habitat to compare restored sites and unrestored sites to a reference ecosystem. After 8 years, diversity and richness differed between the sites, with Forest and Ruderal species more prominent in unrestored sites, and Peatland, Forest, and Wetland species dominant in restored sites. The PRC revealed that the restored site became rapidly dominated by typical peatland plants, the main drivers of temporal changes being Sphagnum rubellum, Pohlia nutans, and Mylia anomala. Some differences remained between the restored and the undisturbed species pools: the former had more herbaceous species associated with wetlands such as Calamagrostis canadensis and Typha latifolia and the latter had more forested species like Kalmia angustifolia throughout the study. PRC revealed to be an efficient tool identifying species driving changes at the community level after restoration. In our case study, examining PRC scores after classifying species according to their preferred habitat allowed to illustrate objectively how restoration promotes target species (associated to peatlands) and how lack of intervention benefits ruderal species.     

Testate Amoeba (Arcellinida,Euglyphida) Ecology along a Poor‐Rich Gradient in Fens of Western Poland

We investigated the relationships between testate amoebae (Arcellinida, Euglyphida), vegetation and water chemistry along environmental gradients in minerotrophic peatlands (fens) in western Poland. We hypothesized that: a) hydrochemistry significantly influences structure of testate amoeba communities, and b) testate amoeba communities are more closely correlated with the hydrochemical variables (environment) than with the vegetation data. Testate amoeba communities and vegetation from 71 sample plots were investigated together with the hydro‐chemistry and hydrology based on 16 environmental variables and vegetation composition. Testate amoeba communities revealed a distinctive poor‐rich gradient in analysed fens. Mineral‐rich habitats, which were dominated by brown mosses, were preferred by a higher number of taxa than acidic habitats, which were dominated by Sphagnum. We recorded a total of 107 testate amoebae taxa. The average species richness of testate amoebae for brown mosses was higher (20) than for Sphagnum (13). We found that testate amoebae communities were similarly correlated with vascular plants, mosses and environmental parameters. Results of direct ordination demonstrate that hydrology, pH, Mg²⁺ and sodium remain the most important environmental control for the entire data set. CCA showed that in case of brown mosses hydrology, sodium and oxygen affect testate amoeba communities significantly whereas in Sphagnum only sodium emerge as most significant determining testate amoeba assemblages. (© 2011 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

One haploid parent contributes 100% of the gene pool for a widespread species in northwest North America

The monoicous peatmoss Sphagnum subnitens has a tripartite distribution that includes disjunct population systems in Europe (including the Azores), northwestern North America and New Zealand. Regional genetic diversity was highest in European S. subnitens but in northwestern North America, a single microsatellite‐based multilocus haploid genotype was detected across 16 sites ranging from Coos County, Oregon, to Kavalga Island in the Western Aleutians (a distance of some 4115 km). Two multilocus haploid genotypes were detected across 14 sites on South Island, New Zealand. The microsatellite‐based regional genetic diversity detected in New Zealand and North American S. subnitens is the lowest reported for any Sphagnum. The low genetic diversity detected in both of these regions most likely resulted from a founder event associated with vegetative propagation and complete selfing, with one founding haploid plant in northwest North America and two in New Zealand. Thus, one plant appears to have contributed 100% of the gene pool for the population systems of S. subnitens occurring in northwest North America, and this is arguably the most genetically uniform group of plants having a widespread distribution yet detected. Although having a distribution spanning 12.5° of latitude and 56° of longitude, there was no evidence of any genetic diversification in S. subnitens in northwest North America. No genetic structure was detected among the three regions, and it appears that European plants of S. subnitens provided the source for New Zealand and northwest North American populations.     

Cranefly (Diptera: Tipuloidea) fauna of a boreal mire system in relation to mire trophic status: implications for conservation and bioassessment

Craneflies (Diptera Tipuloidea) are a typical but poorly known insect group in various moist environments, such as mires. The area of natural mires has strongly decreased in Finland, and there is an urgent need to study and describe the fauna of mires and to determine whether different mire categories support different assemblages of craneflies that might have indicator value. Craneflies were studied using Malaise traps in the Kauhaneva mire system in minerotrophic and ombrotrophic sites, the former subdivided into meso- and oligotrophic sites. A total of 29 cranefly species were recorded. Species richness was highest in mesotrophic sites while the number of species was equally low in oligo- and ombrotrophic sites. Phylidorea squalens, Erioptera flavata, Pedicia rivosa and Tricyphona immaculata were identified as indicators for mesotrophic sites, but no indicators were found for oligo- or ombrotrophic sites. No differences between the species composition of minerotrophic (meso- and oligotrophic combined) and ombrotrophic sites were detected, but when three classes of trophic status were compared, a statistical difference was found. Cranefly species richness in Kauhaneva was low compared to pristine spring habitats. Our results imply, that a focus towards conservation and restoration of mire types with high trophic status would benefit also the conservation of cranefly diversity in the boreal ecoregion. Bioassesments and ecological surveys of craneflies should be designed to cover adequately all trophic status classes within a mire, and especially the mire types with highest trophic status. We also review the distribution and ecology of some potentially regionally threatened cranefly species.     

Allopolyploidy and homoploid hybridization in the Sphagnum subsecundum complex (Sphagnaceae: Bryophyta)

Several complexes of species in Sphagnum (peat mosses) originated through hybridization and allopolyploidy, suggesting that these processes have played a major evolutionary role in this genus. The Sphagnum subsecundum complex includes gametophytically haploid and diploid species in North America. Analyses of 12 microsatellite loci and sequences from two plastid DNA markers show that the evolutionary history of this group is substantially more complex than previously thought. Two taxonomic species, Sphagnum lescurii and Sphagnum inundatum, include both haploid and diploid populations. Within each ploidal level, S. lescurii and S. inundatum are not genetically differentiated. The diploid taxa show patterns of fixed heterozygosity for the microsatellite markers, consistent with an allopolyploid origin. Diploid S. lescurii is an allopolyploid between haploid S. lescurii and (haploid) S. subsecundum. Sphagnum carolinianum is an allopolyploid between haploid S. lescurii and an unknown parent. We detected homoploid hybridization between the haploids Sphagnum contortum and S. subsecundum. Finally, we report three samples of diploid Sphagnum platyphyllum (otherwise haploid) that have an allopolyploid origin involving north‐eastern haploid S. platyphyllum and an unidentified taxon. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 135–151.     

Diversity and Antagonistic Potential of Bacteria Associated with Bryophytes from Nutrient-Poor Habitats of the Baltic Sea Coast

Very little is known about the interaction of bryophytes with bacteria. Therefore, we analyzed bacteria associated with three bryophyte species, Tortula ruralis, Aulacomnium palustre, and Sphagnum rubellum, which represent typical moss species of three nutrient-poor plant communities at the southern Baltic Sea coast in Germany. By use of two cultivation-independent techniques, denaturing gradient gel electrophoresis and single-strand conformation polymorphism analysis of the 16S ribosomal DNA, a high degree of moss specificity was found for associated bacterial communities. This specificity could be further evidenced by a cultivation-dependent approach for the following parameters: (i) plate counts of bacteria on R2A medium, (ii) proportion of antagonistic isolates, (iii) antagonistic activity as well as spectrum against pathogens, and (iv) diversity and richness of antagonistic isolates. The proportion of isolates with antagonistic activity against the pathogenic model fungus Verticillium dahliae was highest for S. rubellum (31%), followed by A. palustre (17%) and T. ruralis (5%). A high percentage (99%) of moss-associated antagonistic bacteria produced antifungal compounds. The high recovery of antagonistic isolates strongly suggests that bryophytes represent an ecological niche which harbors a diverse and hitherto largely uncharacterized microbial population with yet unknown and untapped potential biotechnological applications, e.g., for biological control of plant pathogens.     

Diversity and abundance of nematode-trapping fungi from decaying litter in terrestrial, freshwater and mangrove habitats

Nematode-trapping fungi are ubiquitous in terrestrial habitats in dung, soils, litter and woody debris and they also occur in freshwater, but only one species has been found in marine habitats. The purpose of this study was therefore to investigate whether nematode-trapping fungi occurred in mangrove habitats. To achieve this we assessed the diversity of nematode-trapping fungi on decaying litter from mangroves, freshwater and terrestrial habitats (22 sites) in Hong Kong. Composite samples (n = 1,320) of decaying litter (wood and leaves) were examined and a total of 31 species of nematode-trapping fungi belonging to four genera, Arthrobotrys, Monacrosporium, and Dactylella were recorded. Twenty-nine species reported in this study are new records for Hong Kong and 16 species are new records from mangrove habitats worldwide. Nematode trapping fungi are therefore present in marine environments. Commonly encountered taxa were Arthrobotrys oligospora and Monacrosporium thaumasium which are abundant in all habitats. A. oligospora, M. thaumasium and Arthrobotrys musiformis were frequent (F > 10%). Twenty-six species were rare (0.16–9.32%). Species richness and diversity was higher in terrestrial than in freshwater and mangrove habitats (ANOVA, P < 0.001). A higher mean diversity was observed on decaying leaves as compared to decaying wood in all habitats (P < 0.001). Based on Shannon diversity index, it was also observed that taxa characterized by adhesive nets were more frequent in all habitats. This can be explained by the fact that these taxa may have a better competitive saprotrophic ability which would allow them to compete favourably in nutrient limited environments. Abiotic factors that could be linked to differences in species diversity between decaying wood and leaves are also discussed.     

Local adaptations in bryophytes revisited: the genetic structure of the calcium‐tolerant peatmoss Sphagnum warnstorfii along geographic and pH gradients

Bryophytes dominate some ecosystems despite their extraordinary sensitivity to habitat quality. Nevertheless, some species behave differently across various regions. The existence of local adaptations is questioned by a high dispersal ability, which is thought to redistribute genetic variability among populations. Although Sphagnum warnstorfii is an important ecosystem engineer in fen peatlands, the causes of its rather wide niche along the pH/calcium gradient are poorly understood. Here, we studied the genetic variability of its global populations, with a detailed focus on the wide pH/calcium gradient in Central Europe. Principal coordinates analysis of 12 polymorphic microsatellite loci revealed a significant gradient coinciding with water pH, but independent of geography; even samples from the same fens were clearly separated along this gradient. However, most of the genetic variations remained unexplained, possibly because of the introgression from phylogenetically allied species. This explanation is supported by the small heterogeneous cluster of samples that appeared when populations morphologically transitional to S. subnites, S. rubellum, or S. russowii were included into the analysis. Alternatively, this unexplained variation might be attributed to a legacy of glacial refugia with recently dissolved ecological and biogeographic consequences. Isolation by distance appeared at the smallest scale only (up to 43 km). Negative spatial correlations occurred more frequently, mainly at long distances (up to 950 km), implying a genetic similarity among samples which are very distant geographically. Our results confirm the high dispersal ability of peatmosses, but simultaneously suggested that their ability to cope with a high pH/calcium level is at least partially determined genetically, perhaps via specific physiological mechanisms or a hummock‐forming ability.     

Genetic structure of coastal and inland populations of <Emphasis Type="Italic">Spergularia media</Emphasis> (L.) C. Presl (Caryophyllaceae) in Central Europe

In Central Europe, salt-influenced habitats are restricted mainly to the coastlines of the North Sea and the Baltic Sea. The few natural inland salt sites suffer from size reduction and loss in biodiversity, while anthropogenic salt sites around potash mining dumps experienced recent and massive de novo colonization by numerous halophytes. Our study aimed to elucidate the general patterns of gene flow among coastal and inland salt habitats. We used amplified fragment length polymorphisms to assess the genetic diversity and genetic structure of 18 European populations of the halophyte Spergularia media (Caryophyllaceae), sampled from inland and coastal salt sites of Germany, the Netherlands, Denmark, Austria, France and Italy. Estimates of genetic diversity on the population level were generally low, especially within inland salt habitats. Analyses of molecular variance showed comparatively strong differentiation among populations. Multivariate ordination (principal coordinate analysis) and a NeighborNet analysis revealed four distinct groups of populations that showed good correspondence to their geographic origins. A Bayesian analysis performed with the program Structure as well as high pairwise Φ _ST values supported the results of the phenetic analyses. The observed patterns of diversity and differentiation among inland populations of S. media are best explained by the isolated nature of suitable salt habitats, with concomitant reduction of gene flow to and among these sites. Our data support the hypothesis that the colonization of anthropogenic salt sites by S. media originated from natural inland habitats.