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Acantholimon is an important component of the subalpine steppe flora in the Irano-Turanian region and the second largest genus of Plumbaginaceae with c. 200 cushion-forming subshrubby species. Because the genus has been poorly represented in previous phylogenetic studies, questions regarding its monophyly, phylogenetic relationships and infrageneric classification have not been addressed in a solid evolutionary framework. We used sequences from the nuclear ribosomal internal transcribed spacers and the plastid trnY(GUA)-trnT(GGU) intergenic spacer for 197 accessions of Acantholimon and nine putatively closely related Asian genera in Limonioideae. Contrary to previous results, Bayesian, maximum likelihood and maximum parsimony analyses show that Acantholimon is not monophyletic unless its limits are extended to include species currently placed in eight of these Asian genera. As circumscribed, the new Acantholimon s.l. is sister to Goniolimon and includes the following genera: Bamiania, Bukiniczia, Chaetolimon, Cephalorhizum, Dictyolimon, Gladiolimon, Popoviolimon and Vassilczenkoa. Our phylogenetic analyses also challenge the existing classification at the generic and infrageneric levels. A maximum likelihood reconstruction of ancestral states for morphological characters illustrates the possible scenarios by which the cushion architecture combined with linear acuminate leaves, also present in other steppic Irano-Turanian elements, were acquired in this group of Plumbaginaceae. Our study shows the importance of extensive taxon sampling for phylogenetic reconstruction of species-rich lineages. [ABSTRACT FROM AUTHOR]

We hypothesized that the spatial pattern of genetic variation in different ecological groups of steppe plant species is not similar. To test this hypothesis, we studied genetic variation of the typical steppe plant Goniolimon speciosum (Plumbaginaceae), which is spread across the whole of the Asian steppe, and compared it with the published data on genetic variation of several mountain-steppe species. To elucidate the phylogenetic position of and genetic structure in G. speciosum, we sequenced the nuclear ribosomal internal transcribed spacer (ITS) and plastid trnH-psbA and trnQ-rps16 regions. Goniolimon speciosum was shown to have originated in Central Asia in the Pliocene. We revealed two genetic groups in G. speciosum: a south-eastern (Dahuria, north-eastern Mongolia, southern shore of the lake Baikal and eastern Tyva) group and a broadly distributed western group. This split of evolutionary lineages is estimated to have occurred in the mid-Pliocene. Some samples from the central part of the species area (riverheads of Yenisei and Ob) formed an intermediate genetic group, where most plants had western ITS ribotypes and southeastern plastid haplotypes. This polyphyletic group could have originated due to multiple secondary contacts and subsequent hybridization events. The geographically structured genetic subdivision of the western lineage, based on the ITS data, indicates multiple northward colonizations of G. speciosum from Central Asia that occurred, according to our estimates, in the early Pleistocene. Thus, the history of steppe vegetation is more complicated than has been previously suggested, basing upon limited taxonomic sampling of steppe plants. Species with different ecological preferences have a different history. [ABSTRACT FROM AUTHOR]

Limonium is a widespread genus of halophytes and taxa found on the Atlantic coast include sexual diploids of the L. ovalifolium complex, agamospermous tetraploids of the L. binervosum complex and the triploid L. algarvense. In this study, we investigated: (1) cytotype distribution and diversity within and among populations in an overlapping region of diploid and polyploid Limonium spp. in south-western Iberia and north-western Morocco; and (2) patterns of geographical parthenogenesis and ecological preferences across a latitudinal gradient on the Atlantic coast. We show here for the first time that L. nydeggeri and L. algarvense are found further south in Morocco than previously reported. Genome size and ploidy estimates showed that the distribution of these species is not random at the overlapping region studied: tetraploid apomicts tend to be found at higher latitudes than the sexual diploids and L. algarvense grows in sympatry at the southern boundaries of the diploids. Natural populations showed a constancy in ploidy in these complexes. However, we report for the first time the occurrence of mixed-ploidy populations of L. ovalifolium s.l., euploid triploids in L. algarvense and aneuploids in the L. binervosum complex. On the Atlantic coasts, L. algarvense followed by L. ovalifolium complexes occur significantly more frequently in thermomediterranean and dry ombrotype habitats than the L. binervosum complex. Significant differences were also observed among taxa in the frequency of occurrences on the most common lithological groups. In conclusion, this work presents the first biogeographical insights for the group based in a coarse-scale analysis of data and it provides evidence of ecological differentiation between the studied Limonium complexes. [ABSTRACT FROM AUTHOR]

We investigated a palynological section from middle Miocene sediments at Eskihisar (south-western Anatolia) to establish biogeographic links of the palynoflora and to infer the palaeoenvironment. Four algal palynomorphs, nine spore taxa, eight gymnosperms, three monocots and 67 dicot pollen types were encountered and investigated using the 'single grain method' that combines light microscopy and scanning electron microscopy. Two pollen zones reflect different phases of basin development. Zonal vegetation remained fairly stable across the section and reflects heterogeneous environments including broad-leaved deciduous forest, subtropical forest and sclerophyllous and semi-evergreen oak forest. Conifers were accessory elements in the broad-leaved deciduous forest communities and replaced these at higher elevations. Some herbaceous taxa (Plumbaginaceae) indicate scattered occurrences of sandy and/or rocky soils. Biogeographic affinities are general Northern Hemisphere, North American and East Asian, as also suggested by the macrofossil record. Only two taxa provide potential biogeographic links with the African flora. This suggests that biome shifts of plant taxa between African subtropical/tropical biomes and Anatolian (western Eurasian) temperate forests and shrublands may have been rare in the middle Miocene. [ABSTRACT FROM AUTHOR]

Background and Aims The genus Limonium (Plumbaginaceae) has long been recognized to have sexual and apomictic (asexual seed formation) modes of reproduction. This study aimed to elucidate phylogeographical patterns and modes of reproduction in diploid and tetraploid Limonium species, namely three putative sexual diploid species with morphological affinities (L. nydeggeri, L. ovalifolium, L. lanceolatum) and three related, probably apomict tetraploid species (L. binervosum, L. dodartii, L. multiflorum). Methods cpDNA diversity and differentiation between natural populations of the species were investigated using two chloroplast sequence regions (trnL intron and trnL-trnF intergenic spacer). Floral heteromorphies, ovule cytoembryological analyses and pollination and crossing tests were performed in representative species of each ploidy group, namely diploid L. ovalifolium and tetraploid L. multiflorum, using plants from greenhouse collections. Key Results and Conclusions Genetic analyses showed that diploid species have a higher haplotype diversity and a higher number of unique (endemic) haplotypes than tetraploid species. Network analysis revealed correlations between cpDNA haplotype distribution and ploidy groups, species groups and geographical origin, and haplotype sharing within and among species with distinct ploidy levels. Reproductive biology analyses showed that diploid L. ovalifolium mainly forms meiotically reduced tetrasporic embryo sacs of Gagea ova, Adoxa and Drusa types. Limonium multiflorum, however, has only unreduced, diplosporic (apomictic) embryo sacs of Rudbeckia type, and autonomous apomictic development seems to occur. Taken together, the findings provide evidence of a pattern of 'geographical parthenogenesis' in which quaternary climatic oscillations appear to be involved in the geographical patterns of coastal diploid and tetraploid Limonium species [ABSTRACT FROM AUTHOR]

Background and Aims Limonium is a well-known example of a group of plants that is taxonomically complex due to certain biological characteristics that hamper species' delineation. The closely related polyploid species Limonium vulgare Mill., L. humile Mill. and L. narbonense Mill. are defined species and can be used for studying patterns of morphological and reproductive variation. The first two taxa are usually found in Atlantic Europe and the third in the Mediterranean region, but a number of intermediate morphological forms may be present alongside typical examples of these species. This study attempts to elucidate morphological, floral and karyological diversity representative of these taxa in the Iberian Peninsula.Methods The extent of morphological differentiation was tested through comparison of 197 specimens from both Portugal and Spain using 17 descriptive morphological characters and 19 diagnostic morphometric characters. Analyses of floral morphisms (heterostyly and pollen–stigma dimorphism) and karyological determinations were also conducted.Key Results and Conclusions Discriminant analysis using morphometric variables reliably assigned individuals in natural populations to their respective groups. In addition, the results provide the first direct evidence that L. narbonense and a new species, Limonium maritimum Caperta, Cortinhas, Paes, Guara, Espírito-Santo and Erben, sp. nov., related to L. vulgare are present on Portuguese coasts. Most of these species are found together in mixed populations, especially L. vulgare and L. narbonense. It is hypothesized that taxonomic biodiversity found in sites where distinct species co-occur facilitates the evolutionary processes of hybridization, introgression and apomixis. This study therefore contributes to the elucidation of the taxonomic diversity in L. vulgare-related species and may also help in implementing future conservation programmes to maintain the evolutionary processes generating biodiversity. [ABSTRACT FROM AUTHOR]

Adaptive responses to past climate change may play an important role in the persistence of high-mountain plants, which are vulnerable to global warming. Armeria caespitosa is a high-mountain plant, endemic to the Iberian Central Range. Differences in abiotic environment along the elevational gradient impose two opposing stress gradients (i.e. water stress and duration of the growth season) on the species. Furthermore, the species is found in two interspersed, contrasting microhabitats (rocky outcrops and dry cryophilic grasslands) that have different effects on plants depending of the elevation. As a result of this, the species shows great among-population variation in many reproductive and vegetative traits. We used a common garden approach to determine whether this phenotypic variation has a genetic basis or is the result of plastic responses shaped by heterogeneous environmental conditions. Plants from the high-elevation edge and dry cryophilic grasslands flowered earlier and produced more viable fruits but were smaller. These results confirm that among-population variation in flowering phenology and reproductive performance traits in A. caespitosa is partially genetically based. The results also show that the stronger selection response in favour of early-flowering individuals in populations at the low-elevation edge did not correspond with the greater proportion of early-flowering individuals. Genetic variability associated with flowering onset may be relevant in coping with the impacts of ongoing global warming. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 384-395. [ABSTRACT FROM AUTHOR]

A species-level taxonomic revision of Atocion and Viscaria, based on a recent phylogenetic study, is presented. Atocion includes six species ( A. armeria, A. compactum, A. lerchenfeldianum, A. reuterianum, A. rupestre and A. scythicinum), and Viscaria includes three species ( V. alpina, V. asterias and V. vulgaris). The highest species diversity is found on the Balkan Peninsula. Species descriptions based on a study of morphological characters and generated using the Prometheus description model are provided and are also available in the Sileneae online database. Complete synonymy, notes on the ecology and geographical distribution of the taxa and an identification key are provided. Lectotypes for nine names ( Lychnis helvetica, L. suecica, Silene armeria var. angustifolia, S. berdaui, S. lituanica, S. compacta, S. orientalis, S. reuteriana and Viscaria media) and a neotype for one name ( S. lerchenfeldiana) are assigned. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173, 194-210. [ABSTRACT FROM AUTHOR]

The sea lavender, Limonium wrightii, has six morphs of flower colour variation. The geographical distribution of flower colour morphs is disjunct; the distribution of the pink flower morph is divided into two subregions, and that of the yellow flower morph intervenes between them. The present study aimed to examine the origin of this apparent distribution pattern of flower colour in L. wrightii. Two main hypotheses (i.e. past dispersal events and phenotypic changes by natural selection and/or stochastic processes) have been proposed to account for the origin of leapfrog distribution patterns. To determine which hypothesis was applicable, we conducted a molecular phylogenetic analysis using sequence variation in chloroplast DNA (three regions of intergenic spacers, trnG- trnfM, trnV- trnM, and psbA-trnH). We sequenced 58 accessions of L. wrightii frin 28 islands in the Ryukyu Archipelago and the Izu-Ogasawara Islands, located south of the Japanese mainland, and 12 accessions of four congeneric species. Within L. wrightii, we obtained four lineages of ten haplotypes. These lineages and haplotypes did not correlate with the different flower colours. These results indicate that the formation processes of populations are complex. The haplotypes of the pink flower morph did not show a sister relationship between the two disjunct subregions, indicating that the disjunct populations of the pink flower morphs are unlikely to share the pink flower colour as a result of common ancestry. We conclude that the observed leapfrog distribution pattern is caused by natural selection and/or stochastic processes. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 97, 709–717. [ABSTRACT FROM AUTHOR]

A synopsis of Acantholimon in Turkey is given on the basis of a revisional study conducted in Turkey by the authors. An account of 52 species, ten subspecies, and 17 varieties is given, including keys to the species, subspecies, and varieties, and a general indication of their distribution, as well as their conservation status in Turkey. Seven new taxa, namely A. ekimii sp. nov., A. capitatum ssp. sivasicum ssp. nov., A. calvertii var. glabrum var. nov., A. huetii var. breviscapum var. nov., A. acerosum ssp. longibracteolorum ssp. nov., A. lycaonicum ssp. cappadocicum ssp. nov., and A. halophilum var. coloratum var. nov., are described, together with a few new combinations. A. wiedemannii, A. iconicum, and A. lycaonicum, described from Turkey in the past, but placed under other species as synonyms in the Flora of Turkey, are recognized once again as good species. A. multiflorum and A. parviflorum are given a species status. In Turkey, the presence of A. latifolium, A. araxanum, A. lepturoides, A. hohenackerii, A. senganense, A. fominii, A. petraeum, and A. tragacanthinum is also confirmed. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 154, 397–419. [ABSTRACT FROM AUTHOR]

The compatible solute hypothesis posits that maintaining osmotic equilibrium under conditions of high salinity requires synthesis of organic compounds, uptake of potassium ions, and partial exclusion of NaCl. To assess whether osmotic adaptation in Limonium latifolium proceeds according to this hypothesis, a comprehensive analysis of solute accumulation during NaCl treatments was conducted. Determination of prevailing inorganic ions and establishment of the metabolic profiles for low Mr organic substances revealed that contrary to the mentioned hypothesis the major contributors to osmolarity were constituted by inorganic solutes. independent of salinity, only 25% of this osmolarity resulted from organic solutes such as Suc and hexoses. Proline (Pro), β-alanine betaine, and choline-O-sulfate were minor contributors to osmolarity. Compatible inositols also occurred, especially chiro-inositol, characterized for the first time in this species, to our knowledge. Principal component analysis showed that only a limited number of metabolic reconfigurations occurred in response to dynamic changes in salinity. Under such conditions only sugars, chiro-inositol, and Pro behave as active osmobalancers. Analysis of metabolic profiles during acclimatization to either mild salinity or nonsaline conditions showed that organic solute accumulation is predominantly controlled by constitutive developmental programs, some of which might be slightly modulated by salinity. Osmolarity provided under such conditions can be sufficient to maintain turgor in salinized seedlings. Compartmental analysis of Pro and β-alanine betaine in leaf tissues demonstrated that these solutes, mainly located in vacuoles under nonsaline conditions, could be partly directed to the cytosol in response to salinization. Thus they did not conform with the predictions of the compatible solute hypothesis. [ABSTRACT FROM AUTHOR]

A new species, Acantholimon evrenii sp. nov. is described and illustrated for the first time in sect. Glumaria from Turkey. The species is confined to B7 Elazığ in East Anatolia where it grows on rocky mountain steppes. The diagnostic morphological characters from closely related species are given, along with a discussion dealing with its taxonomic relationship. Its conservation status is indicated. A revised key is also provided for the Acantholimon species with heterophyllous leaves, two or more flowered and four or more bracteated spikelets. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society, 2005, 149, 351–356. [ABSTRACT FROM AUTHOR]

Limonium Mill. sect. Pteroclados (Boiss.) Bokhari (Plumbaginaceae Juss.) was surveyed in this study. Morphology was used for investigating taxonomy as well as for making a cladistic analysis containing 18 ingroup taxa. The two previously recognized subsections Nobiles and Odontolepidae were found to constitute two sister groups. Limonium mouretii (Pit.) Maire shares synapomorphies with both subsections and is positioned as sister group of Limonium sinuatum (L.) Mill. s.l. plus L. lobatum (L. f.) Kuntze but with weak support. Based on the results of the cladistic analyses, a single introduction of a common ancestor to Limonium sect. Pteroclados subsect. Nobiles into the Canary Islands is postulated. Comparisons with cladistic and biogeographical studies of other Macaronesian flowering plant groups were made. An identification key to the taxa, descriptions and discussions on the taxonomy is provided. Morphological differences were found to be minor between Limonium sinuatum, L. bonduellei (T. Lestib.) Kuntze, and L. beaumierianum (Maire) Maire and they are hence suggested to be recognized as subspecies under L. sinuatum. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 144, 461–482. [ABSTRACT FROM AUTHOR]

Three new species, Acantholimon bashkaleicum sp. nov., Acantholimon artosense sp. nov. and Acantholimon hoshapicum sp. nov. (Plumbaginaceae) are described and illustrated. The species are all confined to B9/B10 Van Province in East Anatolia where they grow either on calcareous mountain slopes or on mountain steppes. Diagnostic morphological characteristics from the closely related species are given, along with a discussion dealing with their taxonomic relationships, and their conservation statuses are pointed out. A revised key to the Acantholimon species with heterophyllous leaves, two or more flowered and four or more bracteated spikelets is also provided. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 144, 497–505. [ABSTRACT FROM AUTHOR]

Focuses on the implication of circular DNA cloning and functional expression of a novel n-methyltransferase in the synthesis of the osmoprotectant beta-alanine betaine. Presence of the osmosprotectant beta-alanine in Plumbaginaceae; Synthesis of limonium latifolium by methylation of beta-ala; Inducement of recombinant yeast.