Chromosome number,morphology, pairing,and DNA values of species and hybrids in the genusFallopia (Polygonaceae)

The taxa in this study have in the past been treated together in sect.Tiniaria Meissner ofPolygonum L. s.l., and more recently the large erect rhizomatous herbaceous perennials have been separated from the twining annuals and perennials as the generaReynoutria andFallopia respectively. These taxa range in ploidy level from diploid to octoploid, with base numbers of both 10 and 11 present. The plants are primarily Asiatic in distribution, although many of the large erect perennials are now naturalized in many parts of Europe. Cytological examination has revealed the presence of a number of previously unknown hybrid taxa in the British Isles. The readiness with which hybridisation occurs between taxa of differing base number and ploidy level, and similarities revealed in chromosome morphology, meiotic pairing and in DNA C-value, suggest to the authors that these two genera are best amalgamated underFallopia.     

The use and limits of AFLP data in the taxonomy of polyploid wild potato species in Solanum series Conicibaccata

Solanum sect. Petota (tuber-bearing wild and cultivated potatoes) are a group of approximately 190 wild species distributed throughout the Americas from the southwestern United States south to Argentina, Chile, and Uruguay. Solanum series Conicibaccata are a group of approximately 40 species within sect. Petota, distributed from central Mexico to central Bolivia, composed of diploids (2n = 2x = 24), tetraploids (2n = 4x = 48) and hexaploids (2n = 6x = 64); the polyploids are thought to be polysomic polyploids. This study initially was designed to address species boundaries of the four Mexican and Central American species of series Conicibaccata with AFLP data with the addition of first germplasm collections of one of these four species, Solanum woodsonii, as a follow-up to prior morphological, chloroplast DNA, and RAPD studies; and additional species of series Conicibaccata from South America. AFLP data from 12 primer combinations (1722 polymorphic bands) are unable to distinguish polyploid species long thought to be distinct. The data suggest a complex reticulate history of the tetraploids or the need for a broad downward reevaluation of the number of species in series Conicibaccata, a trend seen in other series of sect. Petota. Separately, through flow cytometry, we report the first ploidy level of S. woodsonii, as tetraploid (2n = 48). The U.S. Government’s right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

The potential role of polyploidy and hybridisation in the further evolution of the highly invasive <Emphasis Type="Italic">Fallopia</Emphasis> taxa in Europe

Japanese knotweed s.l. comprises Fallopia japonica, F. sachalinensis, F. × bohemica and any F2s or backcrosses. The parental taxa were introduced from the East to the West as garden ornamentals in the nineteenth century, and soon spread beyond the confines of the garden to become widespread and persistent weeds. Since only female F. japonica var. japonica was introduced, its impressive spread has occurred solely by vegetative means. However, the initial lack of genetic variability has been complemented by an extensive series of hybridisations in the adventive range. We examine the history, spread, reproductive biology and ecological impact of these species in the West. The role and importance of polyploidy and hybridisation in their invasion of the West is discussed, as are the implications of these factors for the potential further evolution of the group.     

The Japanese knotweed invasion viewed as a vast unintentional hybridisation experiment

Chromosome counts of plants grown from open-pollinated seed from Japanese knotweed around the world have revealed the presence of extensive hybridisation with both native and other introduced taxa. These hybrids fit into three categories: inter- and intraspecific hybrids involving the taxa of Fallopia section Reynoutria (giant knotweeds), hybrids between Japanese knotweed and F. baldschuanica (Regel) Holub and hybrids between Japanese knotweed and the Australasian endemics of the genus Muehlenbeckia. In this minireview, the viability of the different classes of hybrid and the potential threats they pose are discussed in the context of recent examples of allopolyploid speciation, which generally involve hybridisation between a native and an alien species. Such wide hybridisations also challenge accepted taxonomic classifications. Japanese knotweed s.l. provides a fascinating example of the interplay between ploidy level, hybridisation and alien plant invasion. The octoploid (2n=88) Fallopia japonica var. japonica (Houtt.) Ronse Decraene is a single female clone throughout much of its adventive range, and provides an ideal system for investigating the potential for wide hybridisation.     

Agrostis mertensii (Poaceae) in hokkaido

Populations of theAgrostis flaccida-mertensii complex in Hokkaido were examined as part of the taxonomic study of this complex, which has a wide range and comprises a polyploid series of 2x–8x (x=7). The principal aim was to clarify the taxonomic status of hexaploids growing in Hokkaido. Field explorations were made in the mountains and on the coasts in Hokkaido, and the materials were gathered from 19 localities. The occurrence of four cytotypes (2x, 4x, 6x, 8x), as well as some triploids and septaploids, of this complex in Hokkaido was confirmed, and the ranges of respective cytotypes were delineated. Morphological examinations showed that diploids, triploids and tetraploids are referable toAgrostis flaccida and octoploids toA. mertensii. The hexaploids of Hokkaido were morphologically different fromA. flaccida and also from the related hexaploid species,A. tateyamensis, distributed in Honshu. The hexaploids were quite similar to octoploidA. mertensii in all the morphological features examined and also in habitat preference. Septaploids were sterile hybrids between hexaploids and octoploids. A conclusion was drawn that the hexaploid plant of Hokkaido should be grouped with the octoploid in one and the same species,Agrostis mertensii Trin.Agrostis mertensii in this amended sense is discussed from a cytogeographical viewpoint.     

Isoenzyme diversity in Reynoutria (Polygonaceae) taxa: escape from sterility by hybridization

The genus Reynoutria is represented by four taxa in the Czech Republic – R. japonica var. japonica and compacta, R. sachalinensis and R. × bohemica. Using isoenzyme analysis, we determined the degree of genotype variability in all taxa and compared clones of R. japonica var. japonica from the Czech Republic with those from Great Britain. While the rarely occurring tetraploid variety R. japonica var. compacta possesses low variability, the octoploid female clone of R. japonica var. japonica is genetically uniform in the 93 clones sampled and belongs to the same genotype that is present in the whole Europe. R. japonica var. japonica can be fertilized by the pollen of tetraploid R. sachalinensis and a hexaploid hybrid R. × bohemica is produced. In R. sachalinensis, 16 genotypes were found in the 50 clones sampled. R. × bohemica is genetically the most diverse taxon in the study area, with 33 genotypes recorded among 88 clones sampled.     

Allozyme variation in Japanese species ofChionographis (Liliaceae)

Allozyme variation was examined in three diploid taxaChionographis japonica var.japonica, var.kurokamiana, andC. koidzumiana and three tetraploid taxaC. japonica var.kurohimensis, ssp.hisauchiana, and ssp.minoensis. Results show thatC. japonica var.kurokamiana is genetically closer toC. koidzumiana than to var.japonica. In the tetraploid taxa, fixed heterozygosities were found at several loci, and this supports the hypothesis that these taxa are allotetraploids. Furthermore, the tetraploid taxa have many unique alleles not found in the diploid taxa. This suggests that sufficient time has passed since the origin of tetraploids for new mutations to have been fixed.     

国产驴蹄草的细胞地理学研究

为探讨国产毛茛科(Ranunculaceae)驴蹄草属(Caltha)两种植物的演化,该文利用传统染色体压片技术和流式细胞术,并结合前人染色体研究结果,对我国驴蹄草23个居群和花葶驴蹄草10个居群进行了细胞学研究。结果表明:驴蹄草是由四倍体(2n=4x=32)、六倍体(2n=6x=48)和八倍体(2n=8x=64)构成的多倍体复合群,花葶驴蹄草具有四倍体(2n=4x=32)和八倍体(2n=8x=64)两种倍性水平。驴蹄草和花葶驴蹄草均是四倍体较为常见,目前尚未见有二倍体报道。由于驴蹄草和花葶驴蹄草大部分居群采自中国青藏高原地区,可能在冰期时存在古二倍体,其适应性较弱,逐渐被其他的倍性取代,这是由于不同细胞型对环境适应性的结果。驴蹄草可能存在两条进化路线:一条是从甘肃到达云南;另一条是从西藏到达云南。前期分子系统学研究显示花葶驴蹄草与驴蹄草的亲缘关系较近,该研究结果中花葶驴蹄草染色体比驴蹄草要小,花葶驴蹄草可能比驴蹄草相对进化。目前花葶驴蹄草只有10个居群,还需进一步增加居群量来解析其演化路线。     

Asexual spread versus sexual reproduction and evolution in Japanese Knotweed s.l. sets the stage for the “Battle of the Clones”

The invasion of NW Europe by a single clone of male-sterile Fallopia japonica var. japonica by purely vegetative reproduction has demonstrated that reproduction by seed and the genetic diversity associated with it may not be an essential feature of plant invasions, at least in the short term. What is less well known is that a significant proportion of the Japanese Knotweed s.l. involved is not F. japonica var. japonica, but the hybrid between it and F. sachalinensis–F. × bohemica. This hybrid is able to backcross to either parent with the potential to replace the missing male F. japonica; by the same process, the hybrid is generating the genetic diversity so conspicuously lacking in F. japonica. In terms of understanding the population structure in a particular country, it is important to be able to identify hybrids and putative back-crosses. We bring together a mixture of published and unpublished information to provide a comprehensive section on morphological and anatomical aids to identification. Regeneration of plants from stem and rhizome fragments varies by taxon, as do responses to control techniques, underlining the importance of correct identification of these plants. In this paper we look closely at this group of taxa, with special emphasis on the role played by hybridization, with an examination of the genetic make up of seedlings produced in the wild or by artificial hybridizations, and the implications that this will have on the future directions of the invasion process.     

Karyological study in the genusTuberaria sect.Scorpioides (Cistaceae): Taxonomic and evolutionary inferences

A polyploid complex ofTuberaria sect.Scorpioides species in S Spain has been studied. We have found the chromosome numbers n = 12 and 18, which probably originated from the extinct base number x = 6 by means of allopolyploidy. Octoploids (n = 24) seem to be the result of different crossings, and fall morphologically withinT. guttata, T. inconspicua, orT. commutata. Backcrosses between octoploids and tetraploids originate new hexaploids (n = 18), which, once stabilized, behave as amphidiploids. Most of these also fall within the traditional concept ofT. guttata, so that the enormous polymorphism of this taxon seems to be justified.     

Karyomorphological study of the Spiraea japonica complex (Rosaceae)

Reported in this paper is a karyomorphological study on one natural population of each of eight varieties in theSpiraea japonica complex. The interphase and mitotic prophase can be classified into the simple chromocenter type and the proximal type, respectively. The metaphase karyotypes of the eight varieties were formulated as follows:S. japonica var.japonica: 2n=18=14m+2sm+2st;S. japonica var.acuta: 2n=18=11m+4sm+3st;S. japonica var.incisa: 2n=18=12m+4sm+2st;S. japonica var.stellaris: 2n=18=15m+1sm+2st;S. japonica var.acuminata: 2n=18=14m+2sm+2st;S. japonica var.ovalifolia: 2n=18=10m+2sm+2st+4t;S. japonica var.glabra: 2n=18=10m+4sm+4st;S. japonica var.fortunei: 2n=36=17m+16sm+3st. Karyomorphological study reveals that the chromosome numbers within the complex are stable, and that karyomorphological divergence between the varieties lies mainly in chromosome size and organization. Based on the karyomorphological data and geographical distribution of the complex, the differentiation pattern as well as evolutionary mechanism of the complex is evaluated.     

Facilitation in multiple life-history stages: evidence for nucleated succession in coastal dunes

Garden experiments focused on vegetative regeneration were carried out with four invasive taxa of the genus Reynoutria (R. japonica var. japonica, R. japonica var. compacta, R. sachalinensis and a hybrid between R. sachalinensis and R. japonica var. japonica, R. ×bohemica). Regeneration ability of stems and rhizomes, timing of shoot emergence and biomass production were studied under the following treatments: laid horizontally on the soil surface; placed upright; buried in the soil; floating in water. Two different soils (sand and garden loam) representing contrasting nutrient levels were applied. Differences were found in the capability and speed of regeneration, as well as in the quality of shoots produced. Regeneration from stems was less efficient than that from rhizomes in all taxa except R. sachalinensis. R. ×bohemica exhibits higher regeneration potential (61%) than all other taxa and can be considered as the most successful taxon of the Czech representatives of the genus Reynoutria in terms of regeneration and establishment of new shoots. High regeneration capacity was also exhibited by R. japonica var. compacta (52%). Other taxa showed generally lower regeneration rates (R. japonica var. japonica 39% and R. sachalinensis 21%), but under some treatments the percentage of regenerated segments was high, too. R. japonica var. japonica rhizomes regenerated successfully in all three soil treatments but not in the water. An opposite pattern was found for its stems: they regenerated well if exposed to water treatment but in the soil, they did not regenerate at all. Particular taxa responded to the soil type in a contrasting way. R. sachalinensis and R. ×bohemica regenerated better in loam while the opposite was true in R. japonica var. japonica. R. japonica var. compacta produced the tallest and R. ×bohemica the heaviest and most robust shoots. It is concluded that rhizomes are more crucial than stems for the spread of knotweeds through fragmentation and clonal growth, suggesting the importance of soil disturbance.     

Morphological and cytotype variation of wild kiwifruit (Actinidia chinensis complex) along an altitudinal and longitudinal gradient in central‐west China

The Actinidia chinensis complex, a group of commercially important fruits (kiwifruit), is a complex of functionally dioecious lianas of variable ploidy. To understand the cytogeography better and to facilitate breeding in this complex, we examined the ploidy and morphological variations in 16 natural populations of A. chinensis var. chinensis and A. chinensis var. deliciosa across an ecogeographical gradient. Four ploidy levels were found, var. chinensis consisting of diploids and tetraploids and var. deliciosa consisting of tetraploids, pentaploids and hexaploids. Hexaploids were centred in the western Yun‐Gui plateau, tetraploids coexisted with hexaploids or diploids in the middle Yun‐Gui plateau and the Wuling‐Xuefeng mountains, and diploids occurred in the eastern Wuling‐Xuefeng mountains and the Hunan foothills. These findings indicate a gradual, clinal transition from hexaploid to diploid across the elevational and longitudinal gradient. The clear geographical segregation of diploids and hexaploids may have arisen from their differential ecological adaptation in response to altitude and climate, whereas the coexistence of cytotypes (2x–4x, 4x–6x and 4x–5x–6x) might be a result of reproductive barriers, with a particular contribution from the postzygotic reproductive isolation between ploidy races. The geographical pattern and morphological variation of cytotypes suggest a hybrid zone between the varieties in the Wuling‐Xuefeng mountains. The differences in cytotypes which have arisen as a result of ecological adaptation, distribution and morphological characteristics will provide important baseline data for the selection of germplasm and the breeding of kiwifruit. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 72–83.     

A CYTOGEOGRAPHIC STUDY OF ERIOPHYLLUM LANATUM (COMPOSITAE,HELENIEAE)

Analysis of 368 plants derived from 239 natural populations showed that this taxonomically perplexing and wide-ranging species-complex consists of diploids (n = 8), tetraploids, hexaploids and octoploids. Microsporocytes were the source of most of the chromosome counts. Meiosis was basically regular. Multivalent formation was uncommon, but 11 % of all the plants examined had one or more full-sized extra chromosomes. The frequency of plants with extra chromosomes varied significantly among the taxa, from 0 (five varieties) to over 20 % (two varieties). Except in one instance, where one population yielded a diploid and a triploid, different ploidy levels were not found in the same population. The frequency of diploid, tetraploid, hexaploid and octoploid populations was, respectively, 71, 22, 4 and 2%. Variety obovatum appears to be exclusively diploid, and var. aphanactis exclusively tetraploid. Diploids and one or more polyploid levels occurred in the other taxa. No correlation was found between polyploidy and geological history, soils, topography or climate, nor were the polyploids more widely distributed than the diploids. Some of the polyploid populations seem to have been derived from inter-varietal hybridizations, but others do not. The complex has a “pillar” structure in which 10 diploid taxa support a three-level polyploid superstructure. The available evidence suggests that the major role of polyploidy here has been to stabilize the products of intra- and inter-varietal hybridizations.     

Taxonomical study ofAster leiophyllus complex (Compositae) in Kanto district,Japan, with special reference to ploidy level

TheAster leiophyllus complex is complicated taxonomically and includes many related taxa representing polyploid series. It has been reported that this group is diversified throughout the Kanto district, central Japan, and there are three taxa endemic to the Kanto district. However, as neither morphological variation range nor distribution pattern is sufficiently investigated, taxonomic confusion sometimes occurs. In this study, morphological variation was examined in relation to the ploidy level, and taxonomic consideration was given. As a result, four morphologically recognized taxa occur in the Kanto district;A. leiophyllus (2x, 6x),A. leiophyllus var.sawadanus (2x, 3x, 4x),A. semiamplexicaulis (2x), andA. sugimotoi (4x, 5x). The distribution ranges ofA. leiophyllus var.sawadanus, A. semiamplexicaulis andA. sugimotoi are small, whileA. leiophyllus is distributed over most of the Kanto district. Morphological and cytological observations indicate that one of the reasons of the taxonomic confusion is the presence of presumed hybridization betweenA. leiophyllus and other taxa.     

Breakdown of distyly in a tetraploid variety of <Emphasis Type="Italic">Ophiorrhiza</Emphasis><Emphasis Type="Italic"> japonica</Emphasis> (Rubiaceae) and its phylogenetic analysis

We examined the floral morph of tetraploid Ophiorrhiza japonica Blume var. amamiana Hatus. and diploid O. japonica var. japonica to elucidate the association of distyly and ploidy levels. Chloroplast DNA phylogeny was reconstructed to determine the number of tetraploidization events and floral morph shifts in O. japonica. All individuals of O. japonica var. amamiana proved to be long-homostylous, whereas O. japonica var. japonica was distylous with typical long- and short-styled flowers. Distyly is related to the ploidy level. The bagging treatment of flowers indicated that O. japonica var. amamiana is self-compatible and potentially automatically self-pollinating. In cpDNA sequencing analysis, no haplotype was shared between the two varieties. The cpDNA haplotype network displayed the monophyly of O. japonica var. amamiana, suggesting a single origin of this variety. Hence, both tetraploidization and the breakdown of distyly to homostyly in O. japonica var. amamiana likely occurred just once. Because O. japonica var. amamiana having the morphological and cytological entity is recognized as a single lineage and clearly separated from O. japonica var. japonica, this variety can be considered to be a distinct species. We therefore propose to raise O. japonica var. amamiana to the rank of species. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Calamagrostis hakonensis (Poaceae): Distribution and differentiation of cytotypes

Chromosome counts were made for a total of 540 collections ofCalamagrostis hakonensis from 66 localities ranging from Kyushu to Hokkaido. Distribution and habitat preferences of the cytotypes involved are described. Sympatric occurrence of tetraploids (=semidiploids, 2n=28) and higher polyploids, as well as that of septaploids and plants at hexaploid and/or octoploid levels, was confirmed in several localities, and mixtures of hexaploids and octoploids within a population were frequently observed. Plants at hexaploid and octoploid levels were the most abundant and widespread. An examination of pollen of the voucher specimens showed that tetraploids (amphimictic) had good pollen, while higher polyploids (apomictic) were generally devoid of pollen and very rarely produced moderately good pollen, the grains of higher polyploids being larger than those of tetraploids. Distributions of tetraploids and higher polyploids were more precisely delineated through an examination of pollen of many herbarium specimens. In relation to the processes by which the complicated internal structure ofC. hakonensis has been established, the following subjects are discussed: infraspecific hybridization between ecotypically differentiated populations, enrichment of variability through occasional sexual reproduction expected in plants with more than 2n=42, persistence of the variants by apomictic reproduction, and probable roles of some extinct taxa.     

A taxonomic study of KoreanSmilacina

Several taxonomic problems of KoreanSmilacina were examined by statistical methods, based on morphological data. Consequently,S. davurica described in Korean flora differs greatly from originalS. davurica, and closely related toS. bicolor morphologically. Therefore, the present authors treated this new taxon as a variety ofS. bicolor, S. bicolor var.flavovirens N. S. Lee et J. Y. Kim var. nov (“Yondusomdae” in Korean). The distribution in South Korea ofS. bicolor var.bicolor is known as only Mt. Chiri, but this species is collected in Mt. Dukyou, also. The bounds of morphological variations ofS. japonica var.japonica, the short plant size and no hairs on the stem, were revealed. The morphological characteristics ofS. japonica var.japonica andS. japonica var.mandshurica is obscure to distinguish. According to the present study,S. japonica var.mandshurica differs fromS. japonica var.japonica in plant size, size of adaxial leaf epidermal cell, ratio of style/ovary length and shape of stigma. The unrecorded taxon in Korean flora,S. robusta, was given a new Korean name as “Kunsomdae.” AlthoughS. japonica var.mandshurica is the variety ofS. japonica var.japonica, it is more closely related toS. robusta than toS. japonica var.japonica.     

A new cytotype of Jacobaea vulgaris (Asteraceae): frequency,morphology and origin

Jacobaea vulgaris subsp. vulgaris (syn. Senecio jacobaea subsp. jacobaea) constitutes an intricate polyploid complex distributed in Europe. Four cytotypes have been reported in this species, three with euploid (diploid, tetraploid and octoploid; 2n=20, 40 and 80) and one with aneuploid (2n=32) chromosome numbers. Here we report that the diploid chromosome number (2n=20) reported from Bulgaria is due to misidentification with Jacobaea aquatica. On the other hand, we have discovered a new, hexaploid (2n=6x=60) cytotype within J. vulgaris subsp. vulgaris using flow cytometry. The new cytotype occurs within four sympatric populations of otherwise tetraploid and octoploid plants in Pannonia (one locality in the eastern Czech Republic and two localities in southwestern Slovakia) and in Podillya (one locality in western Ukraine). The frequency of hexaploid individuals within 76 studied populations is very low (only 10 of 693 analysed plants), and hexaploids probably represent hybrids between tetraploid and octoploid plants. Three mixed populations with hexaploid plants were subjected to detailed morphological and pollen fertility analyses. Multivariate morphometric analysis reveals partial separation of tetraploid and octoploid plants, whereas hexaploid individuals are similar in morphology to octoploids. In comparison with tetraploids, octoploids and hexaploids exhibit slightly longer ray florets, involucral bracts and tubular florets and more hairy outer achenes. Hexaploid plants display larger pollen grains and lower pollen fertility compared to tetraploids and octoploids.