Conservation of genetic diversity in British populations of the diploid endemic Coincya monensis ssp monensis (Isle of Man Cabbage): the risk of hybridisation with the tetraploid alien, Coincya monensis ssp cheiranthos

Coincya monensis is represented in the British flora by two, cytologically distinct subspecies. Coincya monensis ssp monensis is an endemic diploid with a coastal sand dune distribution that includes a number of isolated populations. Coincya monensis ssp cheiranthos is a tetraploid alien, well established in South Wales in early successional habitats. Both subspecies share similar life form traits, flowering times and pollinators. Cluster analysis and phylogenetic reconstruction based on sequences of the mitochondrial nad4 gene confirmed the distinction between alien and endemic taxa. Tetraploid populations carry more polymorphic RAPDs loci and their genetic diversity is partitioned more within than among populations. In contrast, C. monensis ssp monensis has a distinct population genetic structure. Analysis of the multilocus genetic data confirmed a structure of genetically isolated, endemic population clusters in Scotland, Arran, the Isle of Man and South Wales. Experimental hybridisation showed the two subspecies are interfertile. Multivariate analysis of RAPDs data resolved hybrids between alien and endemic clusters and hybrids contained a proportion of alien-specific polymorphic loci. Hybrids of alien maternal parentage contained the mitochondrial nad4 sequence characteristic of the alien subspecies. Since the alien subspecies can invade mobile sand dune communities from urban sites and compete for pollinators, there is a risk that alien and endemic populations will mix and introgress. Conservation of endemic genetic diversity in Britain will require protection for all C. monensis ssp monensis populations. Currently, the most disjunct endemic population in South Wales is most at risk from introgression.     

EVOLUTION OF THE PYGMY-FOREST EDAPHIC SUBSPECIES OF PINUS CONTORT A ACROSS AN ECOLOGICAL STAIRCASE

Patterns of allozyme variation within and between two of the subspecies of Pinus contorta were examined for the evolutionary relationship between them. In coastal northern California, these subspecies are parapatric. Pinus contorta ssp. contorta occurs on grassy coastal bluffs on the lowest and youngest of a sequence of five marine terraces; P. contorta ssp. bolanderi is endemic to a pygmy forest ecosystem that occurs on the increasingly older and harsher soils of the third, fourth, and fifth terraces. The soils of the upper three terraces are characterized by extreme podzolization, low pH, low nutrient availability, summer drought (with periodic fires), and winter surface flooding above the hardpan. Dune and cliff soils support a tall redwood and Douglas-fir forest between the terraces. Analyses of seeds collected from 11 pygmy-forest and 6 coastal populations showed ssp. bolanderi to have significantly less allozyme variation than spp. contorta. The two subspecies did not show the phylogenetic dichotomy in allozyme allelic constitutions expected for subspecific classification. Within ssp. bolanderi, the pattern of genetic distances correlated better with edaphic differences among sites than with geographic distance. It appears that ssp. bolanderi is a recently evolved derivative of ssp. contorta, and that the low degree of allozyme differentiation among the bolanderi populations may be due to colonization of the sites by small numbers of individuals, or to hitchhiking of allozyme loci linked to loci undergoing strong selection imposed by the severe edaphic conditions typical of bolanderi sites.     

Phylogeography of the white-throated dipper Cinclus cinclus in Europe

Cytochrome b gene sequence variation (902 bp) was used to investigate phylogenetic relationships and genetic differentiation among the four subspecies of European white-throated dipper Cinclus cinclus (Aves; Cinclidae). Six cytochrome b haplotypes were revealed, the most common one (H 13–4) was shared by all seven populations studied. Central European C. c. aquaticus was genetically more diverse than the other subspecies (although not significantly so, due to small sample sizes), whereas our sample of the nominate subspecies from the French Pyrenees showed no haplotype variation. Pairwise comparisons of the four subspecies showed significant genetic differentiation between the taxa except for a lack of differentiation of C. c. hibernicus versus C. c. aquaticus and C. c. cinclus. The genetic structure and the phylogeographic pattern were not related to the geographic distribution of the four subspecies thus disputing the validity of the current infraspecific taxonomy of C. cinclus at least in Europe.Béatrice Lauga and Christine Cagnon contributed equally to this paper.     

Two ecologically distinct subspecies ofHypochaeris radicatal L.

Summary The present paper is part of a series of papers comparing two ecologically distinct subspecies ofHypochaeris radicata L.Hypochaeris radicata ssp.radicata grows in more fertile soils thanH. radicata ssp.ericetorum.The dry-weight yield of plants grown from seeds was measured after a 25 days culture period on varying levels of nitrate. Roots and shoots were analyzed for total organic nitrogen and free nitrate. Dry-weight yield showed an almost identical response over the whole range of nitrate concentrations used. The chemical analysis did not reveal differences between the subspecies that could explain the distribution in the field. In a second experiment the growth rate of the two subspecies was measured under optimal nitrate supply during a two month period. Both subspecies exhibited initially the same relative growth rate but one month after sowing the growth rate of both subspecies declined, with a sharper decline in ssp.ericetorum resulting in an 82 per cent difference in dry-weight between both subspecies two months after sowing. Some pronounced formative differences became evident. Shoot to root ratio of ssp.ericetorum showed little response to increasing nitrate concentrations whereas ssp.radicata was clearly responsive in this respect.The contribution of the tap root in total plant dry-weight was relatively higher in ssp.Ericetorum. The decline in growth rate and the formative differences can explain the absence of ssp.ericetorum from nutrient rich soils. when grown on ammonium ssp.radicata had a slightly but not significantly higher yield than ssp.ericetorum. The difference was even greater at low pH. It is concluded that both subspecies are adapted to ammonium nutrition at pH 4.5 as well as at pH 6 and that the absence of ssp.radicata from acidic soils cannot be explained by the low pH. The possibility that both subspecies have different abilities to reduce their growth when nitrate resources become limiting, and yet remain capable for further growth and reproduction is discussed.Grassland species research group, publ. no.16.     

Variation of plastid and mitochondrial DNAs in the genus Hedysarum

Summary Plastid and mitochondrial DNAs from Hedysarum species of the western Mediterranean basin, H. spinosissimum ssp eu-spinosissimum, H. spinosissimum ssp capitatum, H. carnosum, H. coronarium and H. flexuosum, were compared by restriction endonuclease fragment analysis. ctDNA fragment patterns for ssp eu-spinosissimum and ssp capitatum were indistinguishable in different enzyme digests. An identical ctDNA variation was found in Hpa II digests with two Sardinian populations of ssp capitatum. Each of the two subspecies was characterized by specific mt DNA patterns with Pst I, Bam HI, Sma I and EcoRI. No variation was detected in populations of different geographical origins for a given subspecies. H. carnosum, H. coronarium and H. flexuosum generated specific ct and mt DNA patterns. Comparison of mitochondrial fragments indicated: — a strong homology between the two subspecies, — a closer homology among the three other diploids, each being closer to the other two than to H. spinosissimum subspecies — as was also the case for the plastid genomes.     

Two ecologically distinct subspecies ofHypochaeris radicata L.

Summary Uptake and assimilation of nitrate by two subspecies ofHypochaeris radicata L. were investigated under laboratory conditions as well as in the field.H. radicata ssp.radicata grows on relatively nitrogen-richer soils thanH. radicata ssp.ericetorum. Attempts were made to relate nitrate uptake and nitrate assimilation in the two subspecies to their different distribution in the field.No differences between the two subspecies with respect to nitrate uptake and nitrate assimilation were observed under laboratory conditions. In plants from the field intact tissue nitrate reductase was higher in ssp.radicata than in ssp.ericetorum. The nitrate reductase activity of both subspecies responded positively to nitrate addition.The significance of nitrate uptake capacity and the level of nitrate reductase for the distribution of plants in the field is here discussed.Grassland species research group, publ. no.15.     

Micromorphological and chemical characterisation of Stachys recta L. subsp. serpentini (Fiori) Arrigoni in comparison to Stachys recta L. subsp. recta (Lamiaceae)

Stachys recta L. is a very polymorphous species in which numerous subspecies were recognised. S. recta L. subsp. serpentini (Fiori) Arrigoni is a typical endemism growing on serpentine soils in northern Apennines and particularly in Tuscany (Italy). In order to contribute to a better knowledge of this plant and to its differentiation with respect to S. recta L. subsp. recta, the micromorphological characters (non-glandular and glandular trichomes) and the essential oil composition of the two subspecies were investigated. Micromorphological characters were studied using scanning and transmission electron microscopy, while light microscopy was used for histochemical observations. Essential oil analysis was carried out by gas chromatography and mass spectrometry.In the two examined taxa, the morphology and distribution of glandular and non-glandular trichomes, and the different essential oil composition, may be considered distinctive characters at subspecies level. This is consistent with the taxonomic classification considering S. recta subsp. serpentini a subordinate taxon of S. recta.     

Genetic diversity inferred from AFLP fingerprinting in populations of Onosma echioides (Boraginaceae) from serpentine and calcareous soils

Abstract

Onosma echioides is a non-obligate serpentinophytic borage occurring discontinuously on calcareous and serpentine outcrops at the northwest limit of its range. Mean concentrations of Ca, Mg and heavy metals in root and shoot samples of eight populations from the two soil types were first determined. Subsequently, the genetic polymorphism of the same accessions was estimated by means of Amplified Fragment Length Polymorphism (AFLP) fingerprinting technique. Root and shoot samples from serpentine outcrops showed higher levels of Ni, Cr and Mg, and lower Ca/Mg ratios compared with those from calcareous soils. Based on 353 polymorphic AFLP bands, the two edaphic groups of populations showed comparable levels of genetic diversity. A remarkable genetic differentiation between populations and a high level of within-population genetic variance were found. Results of Mantel's test supported a significant correlation between genetic and geographical distances, while no difference in relation to the edaphic factor was detected. Molecular data suggested isolation as the key factor shaping the infraspecific genetic structure of O. echioides, which may be in relation with the short-distance, zoochorous systems of seed dispersal and pollination of this species.     

膜叶铁角蕨属的形态系统学研究

膜叶铁角蕨属隶属于铁角蕨科,全世界约有30种,我国分布有18种,是该属植物的分布中心之一。到目前为止,膜叶铁角蕨属的物种数目和物种分类还存在很大争议,一些物种的界限和定义还模糊不清,为了得到一个自然的膜叶铁角蕨属分类系统,还需要对膜叶铁角蕨属的物种分类做进一步研究。该文在前人研究的基础上,对膜叶铁角蕨属10种植物的形态特征,包括孢子形态特征、叶柄和根状茎上的鳞片形态特征、叶片形态、羽片形状以及叶脉特征等进行详细观察分析,探讨了各个形态特征间的关系以及膜叶铁角蕨属植物的物种分类。结果表明:膜叶铁角蕨属植物的叶片及羽片等形态特征存在很大区别,叶柄和根状茎上的鳞片以及孢子形态的种间差异虽然不大,但其在大小、形状、颜色等方面的细微差别仍可作为部分种类的鉴定依据。该研究结果为膜叶铁角蕨属植物的物种分类及进一步研究提供了重要依据。     

Patterns of introduction and adaptation during the invasion of Aegilops triuncialis (Poaceae) into Californian serpentine soils

Multiple introductions can play a prominent role in explaining the success of biological invasions. One often cited mechanism is that multiple introductions of invasive species prevent genetic bottlenecks by parallel introductions of several distinct genotypes that, in turn, provide heritable variation necessary for local adaptation. Here, we show that the invasion of Aegilops triuncialis into California, USA, involved multiple introductions that may have facilitated invasion into serpentine habitats. Using microsatellite markers, we compared the polymorphism and genetic structure of populations of Ae. triuncialis invading serpentine soils in California to that of accessions from its native range. In a glasshouse study, we also compared phenotypic variation in phenological and fitness traits between invasive and native populations grown on loam soil and under serpentine edaphic conditions. Molecular analysis of invasive populations revealed that Californian populations cluster into three independent introductions (i.e. invasive lineages). Our glasshouse common garden experiment found that all Californian populations exhibited higher fitness under serpentine conditions. However, the three invasive lineages appear to represent independent pathways of adaptation to serpentine soil. Our results suggest that the rapid invasion of serpentine habitats in California may have been facilitated by the existence of colonizing Eurasian genotypes pre‐adapted to serpentine soils.     

Serpentine and non-serpentine Silene dioica plants do not differ in nickel tolerance

Serpentine soils are hostile to plant life. They are dry, contain high concentrations of nickel and have an unfavorable calcium/magnesium ratio. The dioecious plant Silene dioica (L.) Clairv. (Caryophyllaceae) is the most common herb on serpentine soils in the Swedish mountains. It also commonly grows on non-serpentine soils in the subalpine and coastal area. I have compared the germination frequency, plant establishment and growth of serpentine and subalpine non-serpentine populations in serpentine soil under greenhouse conditions. Further more I have studied the specific effect of nickel on root and shoot growth of serpentine and non-serpentine plants from the subalpine and coastal area in solutions with different concentrations of nickel. Plants from serpentine and non-serpentine populations grew well and in a similar fashion in serpentine soil. Moreover, S. dioica plants, irrespective of original habitat, tolerated enhanced concentrations of nickel when grown in solutions. An analysis of metal content in serpentine plants from natural populations shows that S. dioica has a higher nickel concentration in the roots than in the shoots. The growth studies show that S. dioica is constitutively adapted to serpentine, and that all populations have the genetic and ecological tolerance to grow on serpentine.     

Genetic Variation in Hippophae rhamnoides ssp. sinensis (Elaeagnaceae) Revealed by RAPD Markers

Hippophae rhamnoides ssp. sinensis is endemic to China, and it is a dioecious, outcrossing plant. Although many studies have been undertaken mainly on its agricultural, nutritional, medical, and ornamental value, little is known about its population genetics. This study uses random amplified polymorphic DNA to investigate the genetic diversity and population genetic structure of 13 natural populations of the subspecies sinensis. Fifteen primers amplified 107 reproducible bands, with 95 (88.79%) being polymorphic. The gene diversity within population was 0.168, considerably lower than that of tree species and most perennial, outcrossing species, but higher than that of annual or short-lived, selfing species. The Gst value showed that 18.3% of the total genetic variation resided among populations, a little lower than that of outcrossing species. The present results are quite similar to those previously reported in another subspecies, H ssp. . rhamnoides rhamnoides. The low genetic differentiation among populations in ssp. sinensis may be attributed to the long-distance dispersal of seeds facilitated by birds, in addition to its characteristics of outcrossing, wind pollination, and widespread distribution. No association between genetic distance and geographical distribution was found. The population relationships revealed by the UPGMA dendrogram parallel this result, in that genetic distance did not increase with geographic separation. This pattern of population differentiation may imply the adaptation of ssp. s populations to the local environment, given that its habitats vary greatly across its distribution.     

Genetic differentiation of metallicolous and non-metallicolous Armeria maritima (Mill.) Willd. taxa (Plumbaginaceae) in Central Europe

This study investigates the genetic differentiation within the Central European Armeria maritima (Mill.) Willd. complex with special reference to the metallicolous populations using AFLP markers. Our sampling comprised all metallicolous (ssp. halleri, hornburgensis, bottendorfensis, eifeliaca, calaminaria), and non-metallicolous taxa (ssp. maritima, elongata, alpina). Geographical and genetic distances between populations were moderately positively correlated. Genetic variability of metallicolous and non-metallicolous populations was not significantly different. Lowland populations were clearly differentiated from the alpine populations. Within the lowland group metallicolous and non-metallicolous populations were not genetically differentiated. All lowland populations show a regional differentiation and close relationships to ssp. elongata. Thus, the metallicolous taxa should not be maintained as subspecies. Likewise, their treatment as varieties of a ssp. halleri s.l. is critical because this taxon cannot be consistently characterized throughout its geographical range and may be an artefact itself. If a taxonomical recognition should be considered necessary it is advisable to treat the microendemics as varieties of ssp. elongata.     

Biometrical variability of foliage and cone characters inCupressus bakeri (Cupressaceae)

Biometrical variability of foliage, cones, and seeds were analyzed in 8 native populations (throughout the range) of a relictual species,Cupressus bakeri Jeps. (Cupressaceae) from California and Oregon. The partition of variation within and among populations for this rare plant was investigated and the justification for subspecies status was reviewed. Morphological characters varied significantly, with more of the variation among populations than at the population level. Multivariate analyses indicated close affinities among populations on serpentine sites suggesting the presence of a serpentine race. However, the high degree of variation among populations did not permit their clear separation into the two subspecies described byWolf (1948). We conclude thatC. bakeri should be treated as a monotypic species.     

Serpentine ecotypic differentiation in a polyploid plant complex: shared tolerance to Mg and Ni stress among di- and tetraploid serpentine populations of <Emphasis Type="Italic">Knautia arvensis</Emphasis> (Dipsacaceae)

Background and aims

Serpentine soils impose limits on plant growth and survival and thus provide an ideal model for studying plant adaptation under environmental stress. Despite the increasing amount of data on serpentine ecotypic differentiation, no study has assessed the potential role of polyploidy. We tested for links between polyploidy and the response to serpentine stress in Knautia arvensis, a diploid-tetraploid, edaphically differentiated complex.

Methods

Variation in growth, biomass yield and tissue Mg and Ni accumulation in response to high Mg and Ni concentrations were experimentally tested using hydroponic cultivation of seedlings from eight populations of different ploidy and edaphic origin.

Results

Regardless of ploidy level, serpentine populations exhibited higher tolerance to both Mg and Ni stress than their non-serpentine counterparts, suggesting an adaptive character of these traits in K. arvensis. The effect of ploidy was rather weak and confined to a slightly better response of serpentine tetraploids to Mg stress and to higher biomass yields in tetraploids from both soil types.

Conclusions

The similar response of diploid and tetraploid serpentine populations to edaphic stress corresponded with their previously described genetic proximity. This suggests that serpentine tolerance might have been transmitted during the local autopolyploid origin of serpentine tetraploids.

     

Analysis of the genetic diversity and relationships among and within species of Hippophae (Elaeagnaceae) based on RAPD markers

RAPD markers were used to assess the genetic diversity and inter- and intra-specific relationships of the genus Hippophae L. and to study the correlation between genetic distances and geographic distances among populations of H. rhamnoides ssp. sinensis. The results analyzed by the percentage of polymorphic loci and Shannon information index indicated that a high level of genetic diversity existed both among and within species of the genus Hippophae. In the UPGMA dendrogram, the species or subspecies were clustered into two main groups but not strictly grouped according to sect. Hippophae and sect. Gyantsensis Lian. The multiple regression analysis and Mantel test both indicated a significant correlation between genetic distance and altitude distance among populations of H. rhamnoides ssp. sinensis, and the cluster analysis suggested that the genetic variation among populations of H. rhamnoides ssp. sinensis was linked to their monophyletic origin. Moreover, some degree of genetic differentiation was found among samples collected at different times.     

Song dialects on the Atlantic islands: goldcrests of the Azores ( Regulus regulus azoricus, R. r. sanctae-mariae, R. r. inermis)

Territorial songs in island populations of songbirds are often highly divergent from those of neighbouring continental relatives. This is shown for the three goldcrest subspecies (Regulus regulus azoricus, R. r. sanctae-mariae and R. r. inermis) endemic on six islands of the Azorean archipelago. All investigated populations display a high intra- and inter-individual acoustic variation. On each island, up to six different song types have been found; and a single male sings up to three types. In contrast, all northwestern European populations of R. r. regulus and R. r. anglorum share only a single song type. In playback experiments, none of 18 tested dialect songs of Azorean goldcrests evoked notable territorial reaction in German and Czech goldcrest males (ssp. regulus). Two differing dialect groups of the goldcrest can be distinguished on the Azores. Populations of the eastern islands, São Miguel and Santa Maria, share common song types which are not found on the islands of the central and western groups. Dialect repertoires on the westernmost islands, Flores, Faial and Pico, are dominated by a different song type. In the geologically younger western crater of São Miguel, both western and eastern song types coexist. Acoustic similarities to a population from neighbouring Terceira suggest the western part of São Miguel as the origin for the westward expansion of R. regulus on the Azores.     

RFLP analysis of the wild potato species,Solanum acaule Bitter (Solanum sect. Petota)

Summary Intraspecific variation of a wild potato species, Solanum acaule Bitt., was analyzed by RFLPs of genomic DNA. One hundred and five accessions were selected throughout the distribution area, including all subspecies, i.e., ssp. albicans (hexaploid), ssp. punae (tetraploid), ssp. acaule (tetraploid) and ssp. aemulans (tetraploid). Twenty-seven low-copy DNA clones (probes) were Southern hybridized with EcoRI, EcoRV, HindIII, and XbaI digests of total DNA of all accessions. In total, 238 RFLPs were detected from 94 enzyme x probe combinations. Among them, 49 RFLPs were specific to ssp. albicans, suggesting that the additional third genome is distinct from its two other genomes. RFLPs between and within subspecies were analyzed by principal component analysis. DNA similarities between subspecies coincided with a former taxonomic treatment in the sense that ssp. albicans is the most distantly related to ssp. acaule and ssp. aemulans is distantly related. Subspecies acaule and ssp. punae were indistinguishable. In addition, RFLPs could be used to distinguish groups within subspecies. Subspecies aemulans, confined to Argentina, was divided into two populations, one from the province of La Rioja and the other from the province of Jujuy. In ssp. acaule, some accessions from the southernmost distribution area were clearly distinguishable, while the others varied continuously, showing a geographical cline from Peru to Argentina.Reference to a specific brand or firm name does not constitute endorsement by the US Department of Agriculture over others of similar nature not mentioned     

Floral structure and pollen morphology of two zinc violets (Viola lutea ssp. calaminaria and V. lutea ssp. westfalica) indicate their taxonomic affinity to Viola lutea

Two zinc violets, the yellow form of the Aachen–Liège area and the blue morph of Blankenrode in western Westphalia, have very restricted occurrence on heavy metal waste heaps. Their taxonomic affinities have not been finally resolved. The flower micromorphological analysis presented here indicates that both zinc violets are closely related to the alpine Viola lutea, in line with our earlier published molecular data, but not with the conclusions of other authors. The zinc violets are classed at the rank of subspecies as V. lutea: ssp. calaminaria for the yellow zinc violet and ssp. westfalica for its blue counterpart. Although the violets examined (V. lutea, V. lutea ssp. calaminaria, V. lutea ssp. westfalica) are closely related, there is no evidence that V. lutea ssp. westfalica is a descendent of V. tricolor. Here we provide the most detailed information on generative organ structure in the four violets studied.