Ecological differentiation of Central EuropeanBolboschoenus taxa and their relationship to plant communities

The taxa of theBolboschoenus maritimus group occurring in Central Europe were studied with respect to morphological differentiation, plant communities and soil conditions. Four morphological types ofBolboschoenus were distinguished, differing in habitat conditions, dispersal and distribution: type 1 —Bolboschoenus maritimus subsp.maritimus with narrow triangular fruits; type 2 —B. maritimus subsp.maritimus with wider triangular fruits, type 3 —B. maritimus subsp.compactus with round convex fruits; and type 4 —B. planiculmis auct. with concave fruits. Strong ecological differentiation of these types was found compared with relatively weak morphological diagnostic features. High discriminant efficiency appeared in the relationship to soil chemistry. Differences were also found in habitat-type spectra of individualBolboschoenus morphological types, which was reflected in their affinities to plant communities. Nomenclature, taxonomic classification, and distribution of these taxa in Europe need thorough revision.     

Bolboschoenus glaucus (Lam.) S.G. Smith, a new species in the flora of the ancient Near East

Taxonomic advancement in the genus Bolboschoenus (Cyperaceae, formerly included in the genus Scirpus) have resulted in the re-classification of the plant previously known as Bolboschoenus maritimus (synonym Scirpus maritimus) into several closely-related but distinct Bolboschoenus species This improved taxonomy is of importance for archaeobotanical investigations of ancient sites within the temperate zones, where this genus frequently occurs, because it allows more precise definitions of the ecological requirements and growing habits of each species. Moreover, it details the distinct morphological and anatomical characteristics of the fruit (nutlets) of each species. Using these new nutlet classification criteria, we re-examined charred archaeological specimens which had previously been identified as B. maritimus (or S. maritimus), from five Near Eastern late Pleistocene and early Holocene village sites: Abu Hureyra, Hallan ?emi, Demirk?y, ?atalh?yük and Aswad. Because three of these sites are located in Anatolia, data on the recent occurrence of Bolboschoenus in Turkey were also investigated. All archaeobotanical specimens were found to be B. glaucus. This species was also found to be the most common Bolboschoenus in present-day Turkey, indicating that it has a long history of occurrence in this region. The environmental, ecological and economic implications of this new information suggest that it is entirely feasible that this plant provided late Pleistocene and Holocene Near Eastern people with a dependable and possibly a staple food source.     

Chromosome numbers within the genusBolboschoenus in Central Europe

Two chromosome numbers n=54, n=55 were found inBolboschoenus plants studied from Central Europe (Czech Republic, Slovakia, Hungary, Poland) and coastal regions of Europe (the Netherlands, Sweden). The number n=55 is typical forB. maritimus subsp.maritimus with narrow fruits and mostly also forB. maritimus subsp.compactus; the number n=54 characterizesB. planiculmis auct. The morphological type ofB. maritimus subsp.maritimus with wide fruits represents a stable taxon occurring in freshwater habitats throughout Europe. Its variation in chromosome numbers (both n=54, n=55) indicates a possible hybrid origin, probably resulting from hybridization betweenB. maritimus subsp.maritimus with narrow fruits andB. planiculmis auct. Spontaneous hybridization betweenBolboschoenus taxa in the regions with mixed populations may explain the origin of the intermediate morphological and anatomical characters of plants from some localities and the deviations in chromosome numbers.     

Achene structure and taxonomy of North American Bolboschoenus (Cyperaceae)

Surface and transverse section achene structures ofBolboschoenus maritimus, B. robustus, B. fluviatilis, B. novae-angliae, B. glaucus, and the putative hybridsB. maritimus × B. rubustus andB. glaucus × B. maritimus from North America are described and illustrated with SEM micrographs. The same three type of pericarp structure reported by Browning and Gordon-Gray (1993) from southern Africa were found except that the “type 2” (B. glaucus) was not matched precisely. Intermediate achene structures and other evidence support both the hybrid origin ofB. novae-angliae (=B. fluviatilis × B. robustus) and hybridization betweenB. glaucus andB. maritimus. The new combinationBolboschoenus novae-angliae is made.     

Growth response of Bolboschoenus maritimus ssp. maritimus and B. maritimus ssp. compactus to different trophic conditions

Bolboschoenus maritimus (L.) Palla (=Scirpus maritimus L.) forms extensive stands in the littoral zone of small fishponds and as a weed in rice and maize fields. Within the species, two subspecies are distinguished: Bolboschoenus maritimus subsp. maritimus, B. maritimus subsp. compactus. They differ in ecology, especially in their relationships with trophic conditions and salinity of habitats. To determine growth response of these two types to different nutrient levels, we compared their seasonal development under experimental cultivation at four controlled nutrient levels. Some differences between the subspecies were found to be stable, regardless of nutrient level, namely greater amount of smaller underground tubers and more extensive rhizome system in subsp. compactus compared to less numerous larger tubers and simpler rhizome system in subsp. maritimus. In response to trophic conditions,the plants of subsp. compactus were more resistant to the conditions of the highest trophic level than those of subsp. maritimus, which were stressed. This demonstrates better adaptability and spreading ability of B. maritimus subsp. compactus at high trophic levels.     

The systematic importance of anatomical data in Gagea (Liliaceae) from the Flora Iranica area

Anatomical features of basal leaves, pedicels and tepals of 22 species of Gagea belonging to four sections (Platyspermum, Plecostigma, Gagea and Didymobulbos) are investigated. Anatomical characters are mapped onto a molecular phylogenetic tree and their evolution is evaluated. The systematic importance of the anatomical characters is discussed. Anatomical characters are more systematically useful in the Irano‐Turanian taxa than in Euro‐Siberian taxa. The presence of collenchyma and/or sclerenchyma and the pentagonal outline of the transverse section of the basal leaf is found in Irano‐Turanian taxa of section Platyspermum and is mostly absent in Euro‐Siberian taxa. A diagnostic key based on combined anatomical characters is provided. The level of variation in anatomical characters is greater in the basal leaf than the pedicel and lowest in the tepals. Convergent evolution in anatomical characters is associated with ecological shifts between sunny, open, dry habitats and closed, humid habitats. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 155–177.     

Competition in microcosm between a clonal plant species (Bolboschoenusmaritimus) and a rare quillwort (Isoetessetacea) from Mediterranean temporary pools of southern France

Bolboschoenus maritimus, a clonal species, is locally invasive in Mediterranean temporary pools where it threatens endangered rare plant species such as Isoetes setacea. The combination of management modifications (grazing) and of the progressive accumulation of fine sediments in the pools contributed to the establishment of competitive perennial plants such as B. maritimus. The competitive advantage of B. maritimus on I. setacea has been studied in controlled conditions. The goal of this experiment was to assess the role of environmental conditions in the output of the competition between Bolboschoenus and Isoetes, notably hydrology and soil richness. For this purpose, Isoetes was cultivated alone (three individuals/pot) and with Bolboschoenus (three individuals of both species). The experiment was run with five replicates on six types of sediment (gradient of richness in sand/silt/clay) combined with three hydrological treatments (flooded, wet. and dry). The competitive advantage of Bolboschoenus was measured as the ratio of the production of Isoetes in mixture versus monoculture. The results showed that Isoetes was always outcompeted by Bolboschoenus. However, the competitive advantage of Bolboschoenus on Isoetes, was more related to hydrology than to soil richness. The competitive advantage of Bolboschoenus was very high in wet and flooded conditions and very low in dry conditions. This situation may lead to the extinction, medium-term, of the populations of I. setacea. The introduction of ovine grazing or of cut back practices in temporary pools could reduce the B. maritimus biomass and help toward the conservation of I. setacea populations. Guest editors: B. Oertli, R. Cereghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

Geographic distribution,habitat characterization,and conservation status of <Emphasis Type="Italic">Bolboschoenus</Emphasis> bulrushes (Cyperaceae) in the Hudson River Estuary,USA

Bulrushes of the genus Bolboschoenus are robust, ecologically important sedges occurring in wetlands, including intertidal marshes and mudflats. Despite their importance and multiple serious threats to their habitats, estuarine Bolboschoenus species remain poorly known. We conducted herbarium and field research in order to document historic and current geographic distributions, characterize the habitats, and assess the conservation status of Bolboschoenus species in the Hudson River Estuary, New York, U.S.A. Three species of Bolboschoenus grow in intertidal zones in the Hudson Estuary. Bolboschoenus fluviatilis occurs in the northern, upstream, and freshwater portion of the estuary with multi-year mean surface salinities of 0.078–2.0 ppt. Bolboschoenus robustus occupies the southernmost, downstream, and brackish to saline portion of the estuary with salinities of 4.9–16 ppt. Bolboschoenus novae-angliae occurs in the slightly to strongly brackish region between the other two species with salinities of 1.8–8.0 ppt. The geographic ranges of B. fluviatilis and B. robustus do not overlap, but B. novae-angliae has short zones of sympatry with each of the other two species. Syntopy of B. novae-angliae with each of the other two species is rare. In the Hudson Estuary, B. fluviatilis is secure, but B. novae-angliae and B. robustus are critically imperiled. Threats to future survival of Bolboschoenus species in the Hudson Estuary include competition from invasive plant species (especially Phragmites australis), eutrophication resulting from excess nutrient pollution, and habitat destruction. Our data and analyses provide critical new information for management of existing environmental problems and planned habitat restoration efforts in the Hudson River Estuary.     

Salsola arbusculiformis and S. laricifolia (Chenopodiaceae) in China

To examine diagnostic characters and test phylogenetic relationships between Salsola arbusculiformis and the closely similar S. laricifolia, characters were examined from previous reports and herbarium specimens, and sequences of nrDNA ITS and cpDNA psbB–psbH were compared for these species as well as other representative taxa of Salsoleae. Our results 1) confirm most of the morphological differences highlighted in previous studies, such as those of leaf size, bracts, bracteoles, style, fruiting perianth, and anther appendages; 2) show that S. arbusculiformis is a typical Irano–Turanian species, mainly distributed from Iran and Turkmenistan to westernmost China (Tacheng, Ili, and Bole regions, Xinjiang), while S. laricifolia is a central Asian floristic element mainly distributed from east Kazakhstan and Kyrgyzstan to northern China (north Xinjiang, inner Mongolia, Ningxia and Gansu provinces), and Mongolia; and 3) strongly support their sister relationship, with samples from five different populations clustering together for each of the two species.     

Environmental gradients shape the genetic structure of two medicinal Salvia species in Jordan

• Environmental gradients, and particularly climatic variables, exert a strong influence on plant distribution and, potentially, population genetic diversity and differentiation. Differences in water availability can cause among‐population variation in ecological processes and can thus interrupt populations’ connectivity and isolate them environmentally. The present study examines the effect of environmental heterogeneity on plant populations due to environmental isolation unrelated to geographic distance.
• Using AFLP markers, we analyzed genetic diversity and differentiation among 12 Salvia spinosa populations and 13 Salvia syriaca populations from three phytogeographical regions (Mediterranean, Irano‐Turanian and Saharo‐Arabian) representing the extent of the species’ geographic range in Jordan. Differences in geographic location and climate were considered in the analyses.
• For both species, flowering phenology varied among populations and regions. Irano‐Turanian and Saharo‐Arabian populations had higher genetic diversity than Mediterranean populations, and genetic diversity increased significantly with increasing temperature. Genetic diversity in Salvia syriaca was affected by population size, while genetic diversity responded to drought in S. spinosa. For both species, high levels of genetic differentiation were found as well as two well‐supported phytogeographical groups of populations, with Mediterranean populations clustering in one group and the Irano‐Turanian and Saharo‐Arabian populations in another. Genetic distance was significantly correlated to environmental distance, but not to geographic distance.
• Our data indicate that populations from moist vs. arid environments are environmentally isolated, where environmental gradients affect their flowering phenology, limit gene flow and shape their genetic structure. We conclude that environmental heterogeneity may act as driver for the observed variation in genetic diversity.

     

Systematic Rearrangement of Korean <Emphasis Type="Italic">Scirpus</Emphasis> L. s.l. (Cyperaceae) as Inferred from Nuclear ITS and Chloroplast <Emphasis Type="Italic">rbcL</Emphasis> Sequences

The genus Scirpus L. s.l. (Cyperaceae) has been accepted as a polyphyletic taxon by most plant taxonomists. This genus was separated into different genera by several different authors: Scirpus s. str., Trichophorum Pers., Bolboschoenus (Asch.) Palla, Schoenoplectus (Rchb.) Palla, and Schoenoplectiella Lye. The heterogeneity of Korean Scirpus s.l. has not yet been studied. We examined 17 taxa of Korean Scirpus s.l. by morphological characters and phylogenetic analyses based on nuclear ITS and chloroplast rbcL sequences. Phylogenetic analyses using maximum parsimony, maximum likelihood, and Bayesian method provided sufficient resolution. The phylogeny revealed the polyphyly of Korean Scirpus s.l. with five distinct clades. These clades correspond to Bolboschoenus, Schoenoplectus s.str., and Schoenoplectiella, Scirpus s.str., and Trichophorum, respectively. These five genera were delimited in terms of the morphology of tuber, bract, and inflorescence. By virtue of our findings, we suggest that the 17 taxa of Korean Scirpus s.l. should be placed into five genera as follows: Bolboschoenus (two species), Schoenoplectus (three species), Schoenoplectiella (six species), Scirpus s.str. (five species), and Trichophorum (one species).     

Local adaptation to different phytogeographic regions: habitat‐related variations in seed germination in response to temperature and salinity for two medicinal Salvia species from Jordan

Salvia spinosa L. and Salvia syriaca L. are perennial medicinal herbs that occur in the Mediterranean, Irano‐Turanian and Saharo‐Arabian phytogeographic regions of Jordan. With respect to the seed germination requirements, prevailing environmental conditions in each phytogeographic region may promote local adaptation and consequently affect the distribution range of the species. Using seeds of both species collected from populations across the three regions, we tested responses to variations in temperature and salinity under laboratory conditions. Both species showed significant differences in cumulative germination percentages and germination rates (modified Timson Index) with temperature, while origin only significantly affected S. spinosa seeds. Both species germinated best under the highest temperature regime (32/20°C). The low temperature regime (8/4°C) completely inhibited germination in S. syriaca, wheras it led to 80% to 95% germination in S. spinosa, with significant variation being recorded between the phytogeographic regions. For both species, salt solutions of 0, 25 and 50 mM NaCl yielded the highest germination percentages and rates, which sharply and significantly declined at higher concentrations (100 and 200 mM NaCl). Our results provide evidence of local adaptation of the study species to salinity and temperature in the respective maternal environments, particularly in the Irano‐Turanian and Saharo‐Arabian regions. Such differentiation should be accounted for in future conservation planning.     

Effect of anatomical structure on the buoyancy of achenes of two subspecies ofBolboschoenus maritimus

Differences between two subspecies ofBolboschoenus maritimus (L.)Palla (=Scirpus maritimus L.) in different habitats (littoral and terrestrial) are partly due to variations in the buoyancy of their achenes, caused by different anatomical structure of the fruits. This hypothesis was tested by comparing fruit buoyancy and anatomy of both subspecies at several localities. Plants were sampled fromBolboschoenus maritimus subsp.maritimus (plants with branched inflorescences and triangular fruits, 4 localities) andB. maritimus subsp.compactus (Hoffm.)Hejný (plants with compact inflorescences and prevailing concave fruits, 4 localities) in the Czech and Slovak Republics, andB. maritimus subsp.compactus (plants with compact inflorescences and convex fruits, 1 locality) from the coast of South Sweden. From each locality a plant with several tubers connected by rhizomes was taken and transferred into the experimental garden; from these cultivated plants (clones) mixed samples of fruits collected from each clone were used for buoyancy tests and a study of their anatomy. Fruit buoyancy was found to depend on the anatomical structure of the pericarp—the presence of aeriferous tissue in the exocarp serves as the main floating organ. This differentiated all plants studied into two groups: short-floating subsp.maritimus, with a thin or none acriferous tissue in the exocarp, and long-floating subsp.compactus with well developed aeriferous tissue. Differences in fruit buoyancy corresponds to differences in the prevailing habitat types of both subspecies studied in the inland sites: the frequent occurrence of subsp.compactus in temporarily flooded terrestrial habitats contrasts with the prevalence of subsp.maritimus in the littoral of water bodies. This is an example of intra-specific ecological differentiation leading to differentiation in wetland communities.     

Efficiency of three halophyte species in removing nutrients from saline water: a pilot study

Saline wetlands may be well suited for purifying contaminated water from saline agriculture and aquaculture or from freshwater-based agriculture in areas subject to increased salinity. However, case studies on the nutrient removal efficiency of halophyte species are scarce, especially for temperate regions. Here we tested the nutrient removal efficiency and ability to store nutrients in aboveground and belowground biomass of three halophyte species, Aster tripolium, Bolboschoenus maritimus subsp. compactus, and Spartina anglica, in a greenhouse microcosm experiment at two salinity levels. Nutrient removal from water differed among the species: Spartina had the highest nitrogen removal, Bolboschoenus and Spartina had the highest phosphorus removal. The species also differed in the allocation of the nutrient uptake. Bolboschoenus had the highest absolute uptake of nitrogen and phosphorus in shoots, whereas Spartina had the highest uptake of nitrogen and phosphorus in roots. The applicability of these three species in constructed saline wetlands depends on the local salinity and water regime.     

The effect of submergence on tuber production and dormancy in two subspecies ofBolboschoenus maritimus

Two subspecies ofBolboschoenus maritimus (L.)Palla were cultivated in submerged and non-submerged conditions. The production of tubers, the proportion of tubers remaining dormant and the time between production and reactivation were investigated. Total tuber production was found to be dependent on subspecies and may be considered a biological character of the subspecies.Bolboschoenus maritimus ssp.compactus, a subspecies of less, flooded habitats, produced more tubers with a greater capacity for growth leaving a smaller proportion of dormant tubers. Inactivity of tubers and their reactivation after one or two years were enhanced by submergence in both subspecies. The proportion of tubers that are permanently dormant as well as the dormancy of the whole plant are important features influencing the survival of populations ofB. maritimus over periods of unfavourable conditions.     

Dionysia assadii sp. nov. (Primulaceae: sect. Dionysiopsis) from southern Iran

The new species Dionysia assadii (Primulaceae) from the Irano–Turanian region in Fars province, Iran, is described and illustrated. It is compared with its closest relative Dionysia esfandiarii Wendelbo and differs from it in leaf shape, leaf margin, hair density of leaves and other organs, shape of calyx and bracts, color of corolla and status of lobes, form and color of seeds, as well as the length of anthers and style in longistylous flowers.     

Field studies on the ecology ofBolboschoenus maritimus (L.)Palla (Scirpus maritimus L. s. l.)

In an ecological field study, plants ofBolboschoenus maritimus (L.)Palla (Scirpus maritimus L. s. l.) growing in oligohaline and in mesohaline soils were compared. Differences between both populations mainly concerning osmotic potential, water potential and ionic concentrations could be related with the salt content at both sites, whereas transpiration and photosynthesis did not differ significantly. Water potential of the cell sap was affected by several external factors.     

Patchy Distributions of Competitors Affect the Growth of a Clonal Plant When the Competitor Density Is High

Environments are patchy in not only abiotic factors but also biotic ones. Many studies have examined effects of spatial heterogeneity in abiotic factors such as light, water and nutrients on the growth of clonal plants, but few have tested those in biotic factors. We conducted a greenhouse experiment to examine how patchy distributions of competitors affect the growth of a rhizomatous wetland plant Bolboschoenus planiculmis and whether such effects depend on the density of the competitors. We grew one ramet of B. planiculmis in the center of each of the experimental boxes without competitors (Schoenoplectus triqueter), with a homogeneous distribution of the competitors of low or high density, and with a patchy distribution of the competitors of low or high density. The presence of competitors markedly decreased the growth (biomass, number of ramets, number of tubers and rhizome length) of the B. planiculmis clones. When the density of the competitors was low, the growth of B. planiculmis did not differ significantly between the competitor patches and competitor-free patches. However, when the density of the competitors was high, the growth of B. planiculmis was significantly higher in the competitor-free patches than in the competitor patches. Therefore, B. planiculmis can respond to patchy distributions of competitors by placing more ramets in competition-free patches when the density of competitors is high, but cannot do so when the density of competitors is low.     

Phytogeography of Genu and Homag,two mountains with an Irano–Turanian flora in the Saharo–Sindian regional zone,south Iran

The Saharo–Sindian regional zone encompasses the flat and arid areas of North Africa, the Arabian Peninsula, southern Iran and the deserts of Pakistan and west India. There are some scattered mountains situated within this area, like Hoggar in Sahara, Saint Catherine in Sinai and Genu and Homag in southern Iran. These highlands serve as interglacial refugia for cold adapted plant species. In the present study, phytogeographical patterns and relationships of the flora of Genu and Homag mountains are described and discussed in relation to the phytogeography of the flora of low‐lying Hormoz Island. According to a chorological assesment of the flora, Genu and Homag mountains belong to the Irano–Turanian region with 59% of the species restricted to this area. In contrast, the surrounding lowland plains are part of the Saharo–Sindian area with a rather high proportion of widespread species (17%) and Somalia–Masai‐linking elements (20%). It is noteworthy that several Turanian enclaves also occur in the lowland zone. Furthermore, the distributional patterns imply that the mountainous Irano–Turkestanian region is an integrated area which is supposedly distinct from the Turanian lowland areas in the north and from the Saharo–Sindian lowland areas in the south. On the other hand, the expansive floras of Turanian and Saharo–Sindian regions are linked to each other. Endemic species in lowland areas in south Iran are mostly either frost sensitive vicariants of cold adapted Turanian species or of Saharo–Sindian origin, while the highland endemics in the area trace their origins to the Irano–Turkestanian region.