Polyploidy and new chromosome counts in Anaphalis (Asteraceae: Gnaphalieae) from the Qinghai‐Tibet Plateau of China

Abstract Anaphalis is one of the largest genera of the Asian Gnaphalieae (Asteraceae) and is most diversified on the Qinghai‐Tibet Plateau. The chromosome numbers and karyomorphology of six species from seven populations were investigated for the first time for this region. Chromosome numbers have been newly documented for four species: Anaphalis deserti (2n = 56 = 24 median [m]+ 24 submedian [sm]+ 8 acrocentric [st]); Anaphalis plicata (2n = 56 = 26m + 30sm); Anaphalis xylorhiza (2n = 28 = 2 median point [M]+ 14m + 12sm); and Anaphalis rhododactyla (2n = 56 = 32m + 24sm). Two further counts are in agreement with the previously reported numbers, that is, Anaphalis royleana (2n = 28 = 4M + 6m + 18sm) and Anaphalis margaritacea (2n = 28 = 12m + 16sm). A new polyploid cytotype (2n = 56 = 26m + 30sm) was found in the Zougong population of A. margaritacea. Polyploidy is for the first time reported for Anaphalis, with four out of seven counts being tetraploid. Our cytological results suggest that polyploids might have played an important role in the evolution of Anaphalis on the Qinghai‐Tibet Plateau.     

Polyploidy and new chromosome counts in Anaphalis(Asteraceae:Gnaphalieae) from the Qinghai-Tibet Plateau of China

Anaphalis is one of the largest genera of the Asian Gnaphalieae (Asteraceae) and is most diversified on the Qinghai-Tibet Plateau. The chromosome numbers and karyomorphology of six species from seven populations were investigated for the first time for this region. Chromosome numbers have been newly documented for four species: Anaphalis deserti (2n = 56 = 24 median [m] + 24 submedian [sm] + 8 acrocentric [st]); Anaphalis plicata (2n = 56 = 26m + 30sm); Anaphalis xylorhiza (2n = 28 = 2 median point [M] + 14m + 12sm); and Anaphalis rhododactyla (2n = 56 = 32m + 24sm). Two further counts are in agreement with the previously reported numbers, that is, Anaphalis royleana (2n = 28 = 4M + 6m + 18sm) and Anaphalis margaritacea (2n = 28 = 12m + 16sm). A new polyploid cytotype (2n = 56 = 26m + 30sm) was found in the Zougong population of A. margaritacea. Polyploidy is for the first time reported for Anaphalis, with four out of seven counts being tetraploid. Our cytological results suggest that polyploids might have played an important role in the evolution of Anaphalis on the Qinghai-Tibet Plateau.     

Phylogenetic relationships in the genus Leontopodium (Asteraceae: Gnaphalieae) based on AFLP data

The genus Leontopodium comprises 30–41 species. The centre of diversity is the Sino‐Himalayan region in south‐western China, where about 15 species occur. The two species native to Europe, L. alpinum (known as the common ‘Edelweiss’) and L. nivale, are part of the cultural heritage of the people living there. Despite its importance, very little is known about the systematics of the genus. Because recent molecular studies have shown that species within this genus are closely related and difficult to distinguish with rDNA and cpDNA data, we used AFLPs to obtain a more detailed understanding of the phylogeny of the genus. Our main aims were as follows: (1) to clarify species relationships within the genus; and (2) to reveal information about the biogeography of the genus. We used AFLPs with six primer combinations to investigate 216 individuals in 38 populations of 16 different species. With AFLPs, we were able to recognize 10 different groups, all of which had strong bootstrap support. These results were also congruent with the morphology‐based taxonomy of the genus. Most private and rare fragments were found in the Yunnan region (south‐western China) relative to Europe and Mongolia/central China, suggesting a long‐lasting in situ history of populations in the centre of diversity of the genus. Our results illustrate the utility of AFLPs to resolve phylogenetic relationships between these closely related species. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 364–377.     

New or rare chromosome counts in the genus Artemisia L. (Asteraceae, Anthemideae) from Armenia and Iran

Nineteen chromosome counts of 12 Artemisia species are reported from Armenia and Iran. Three of them are new reports, sewn are not consistent with previous counts and the remaining are confirmations of very scarce (one to three) previous data. Two basic chromosome numbers (x = 8 and 9) were found, each with several ploidy levels. Chromosome number reduction arising from fusion homozygosity was noted, confirming earlier studies.     

Chromosome counts in Asian Artemisia L. (Asteraceae) species: from diploids to the first report of the highest polyploid in the genus

This work comprises 24 reports of chromosome numbers in 24 Artemisia L. species from Asia. Ten are included in the subgenus Dracunculus Besser and the rest belong to other subgenera. Seven counts are new reports, 14 are consistent with scarce previous ones, and three contribute new ploidy levels. That carried out in A. medioxima reports the highest ploidy level ever counted for the genus (16 x ). There is only one species with x  = 8 as the basic chromosome number. In the remaining x  = 9-based species, ploidy levels range from 2 x to 16 x , illustrating the great role played by polyploidy in the evolution of the genus.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 301–310.     

Chromosome numbers in the tribe Anthemideae (Asteraceae) from north-east Anatolia

Twenty-two chromosome counts of 19 taxa in the tribe Anthemideae of the family Asteraceae are reported from north-east Anatolia, Turkey. The taxa belong to the subtribes Achilleinae (four Achillea taxa), Anthemidinae (five Anthemis taxa), Artemisiinae (two Artemisia species), Leucantheminae (one Lecanthemum species), Matricariinae (two Tripleurospermum species) and Tanacetinae (five Tanacetum taxa). Six counts are new reports, seven are not consistent with previous counts, and the remainder are confirmations of very limited previous data. Most of the populations of Anthemideae studied have the basic number x  = 9, with ploidy levels ranging from 2 x to 7 to 8 x , but dysploidy is also present, with one diploid species, Artemisia austriaca , having x  = 8.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 203–211.     

Chromosome numbers and meiotic studies in species of Senecio (Asteraceae) from Argentina

Meiotic chromosome counts, chromosomal behaviour and meiotic configurations of ten taxa of Senecio from Argentina were examined. Most counts are original: S. crepidifolius DC., S. francisci Phil. and S. octolepis Griseb. var. saltensis (Hicken) Cabrera & Zardini have 2 n  = 40, and S. chrysolepis Phil., 2 n  = 80 + 8B. We confirmed previous reports for S. deferens Griseb. (2 n  = 40 + 4B), S. filaginoides DC. var. filaginoides (2 n  = 40), S. hieronymi Griseb. (2 n  = 40 + 7B), S. pampeanus Cabrera (2 n  = 40) and S. rudbeckiifolius Meyen & Walp. (2 n  = 40 + 7B). In S. bracteolatus Hook. & Arn. var. bracteolatus , we found a new number (2 n  = 40) that differs from the one cited previously. In four species, numerical polymorphisms for B-chromosomes were observed. Several of the analysed species presented secondary bivalent association. This phenomenon, together with other evidence, supports x  = 5 as the basic chromosome number. The number of chiasmata and their positions were also surveyed, with the finding that open bivalents were the most frequent meiotic figures and terminal chiasmata the preferential position. These features are related to recombination rate, and the success and persistence of these polyploids. We discuss some systematic and evolutionary aspects in the light of cytogenetic data and conclude that polyploidy is the major force modelling the chromosome evolution within this genus.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 465–474.     

Contribution to the genome size knowledge of New World species from the Heliantheae alliance (Asteraceae)

This article contributes first genome size assessments by flow cytometry for 16 species, 12 genera, and 3 tribes from family Asteraceae, mostly belonging to the Heliantheae alliance, an assembly of 13 tribes from subfamily Asteroideae with a large majority of its species in the New World. Most genome sizes are accompanied by their own chromosome counts, confirming in most cases, although not all, previous counts for the species, and revealing possible cases of unknown dysploidy or polyploidy for certain taxa. The data contribute to the pool of knowledge on genome size and chromosome numbers in the family Asteraceae and will further allow deeper studies and a better understanding on the role of dysploidy in the evolution of the Heliantheae alliance. However, we still lack data for tribes Chaenactideae, Neurolaeneae, Polymnieae, and Feddeeae (the latter, monospecific) to complete the alliance representation.     

New or rare chromosome counts in the genera Artemisia L. and Mausolea Bunge (Asteraceae, Anthemideae) from Uzbekistan

Seventeen chromosome counts of 14 Artemisia and one Mausolea species are reported from Uzbekistan. Five of them are new reports, two are not consistent with previous counts and the remaining are confirmations of very scarce (one to four) previous data. Two basic chromosome numbers (x = 8 and x = 9) and two ploidy levels (2x and 4x) were found. Some correlations between ploidy level, morphological characters and distribution are noted.     

Cytogeography of European perennial species of Cyanus (Asteraceae)

To reveal the general cytogeographical pattern of Cyanus section Protocyanus in Europe, DNA ploidy and/or chromosome numbers were newly examined for 160 populations by flow cytometry (450 plants) and/or chromosome counting (30 plants). Furthermore, previously published karyological data were revised (236 records). Our analyses confirmed chromosome counts of 2n = 22 for all newly investigated samples of the C. triumfetti group (the records for C. semidecurrens and C. ternopoliensis are new), C. diospolitanus and C. achtarovii; 2n = 44 for C. montanus and C. mollis; and 2n = 20 for C. lingulatus, C. napulifer, C. nissanus, C. orbelicus, C. thirkei, C. tuberosus and C. velenovskyi. The chromosome count of 2n = 20 is the first report for C. epirotus. The cytotype 2n = 40 was newly recorded for the Crimean endemic C. fuscomarginatus and Calabrian and Greek populations of C. graminifolius. The cytotypes 2n = 20 and 2n = 40 were confirmed for C. pindicola. For the first time triploidy (2n～3x～30) was found in C. nissanus, C. thirkei and in a newly discovered hybrid, C. epirotus × C. graminifolius. Two contrasting ecogeographical patterns emerged: cytotypes derived from the base chromosome number x = 11 (2n = 22, 44) are widespread in northern latitudes and ecologically diverse, whereas cytotypes with x = 10 (2n = 20, 30, 40) are confined to mountains in southern Europe. In general, tetraploids have smaller ranges than diploids. The new combinations Cyanus section Protocyanus (Dobrocz.) Ol?avská comb. nov. and Cyanus ternopoliensis (Dobrocz.) Ol?avská comb. nov. are provided. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 230–257.     

Chromosome numbers in the tribes Anthemideae and Inuleae (Asteraceae)

Twenty-two chromosome counts of 19 taxa (21 populations) in the tribe Anthemideae and one member (one population) of the tribe Inuleae of the family Asteraceae are reported. The Anthemideae studied belong to the subtribes Artemisiinae (14 Artemisia taxa, and one species each of the genera Dendranthema , Filifolium and Neopallasia ) and Tanacetinae (one species each of the genera Lepidolopha and Tanacetopsis ). From the Inuleae, we studied one Inula species. Five counts are new reports (including two at generic level), six are not consistent with previous counts and the remainder are confirmations of very limited (one to four records) previous data. Most of populations of Anthemideae studied have the basic chromosome number x  = 9, with ploidy levels ranging from 2 x to 10 x . Dysploidy is also present, with two x  = 8 diploid taxa. The species of Inuleae studied is a diploid with x  = 10, also indicating dysploidy, other members of the same genus Inula having basic numbers of x  = 9 or 8.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 77–85.     

Contribution to the karyological knowledge of Echinops (Asteraceae, Cardueae) and related genera

Twenty-six chromosome counts were made of several genera of the tribe Cardueae from various European and Asian provenances: Acantholepis (one species studied), Amphoricarpus (1), Chardinia (1), Echinops (14 species, 15 populations), Siebera (1), Staehelina (3) and Xeranthemum (4). Eleven of the reports are made for the first time, ten confirm previous counts, while the remainder report disparities with earlier records. The existence of different basic chromosome numbers and ploidy levels suggests dysploidy and polyploidy as the main mechanisms of chromosomal evolution in the taxa considered.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 337−344.     

Hybridization between diploid Centaurea pseudophrygia and tetraploid C. jacea (Asteraceae): the role of mixed pollination,unreduced gametes,and mentor effects

We studied hybridization between the diploid Centaurea pseudophrygia and the tetraploid C. jacea by performing crossing experiments and screening natural populations using flow cytometry. The experiments confirm that the studied species exhibit strong reproductive isolation. Interspecific hybrids were formed at a low frequency, including triploids (originating from reduced gametes) and tetraploids (involving unreduced gametes of the diploids). In contrast, hybrids were almost absent among seeds and adult plants of natural mixed populations and among the offspring from experimental pollinations with a mixture of pollen of both ploidy levels. We found that mixed pollination is an important mechanism for preventing hybridization between plants of different ploidy levels and sustaining the reproduction of the tetraploids. A mentor effect (induced selfing in the presence of pollen of different ploidy levels) was observed in both diploids and tetraploids, reinforcing the reproductive isolation between cytotypes. Higher ploidy levels (pentaploid, hexaploid) involving unreduced gametes of the tetraploid species were identified. Notably, pentaploids were discovered for the first time in Centaurea sect. Jacea. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 93–106.     

Karyotype evolution and cytotaxonomy in Brazilian species of Rhynchospora Vahl (Cyperaceae)

Karyotypes were analysed in 113 populations of 34 species of Rhynchospora (Cyperaceae). They showed typical holocentric chromosomes lacking primary constrictions. Nucleolar constrictions were observed in the terminal regions of a few chromosomes (except in R. nanuzae , in which they were interstitial). The chromosome numbers varied between 2 = 4 and 2n = 50. The primary base number seems to be x = 5; x = 6 and x = 9 are secondary. Dysploidy occurred at low frequency, whereas polyploidy seemed to be the predominant event in the karyotype evolution of the genus. Karyotypic differences were found in R. consanguinea var. speciosa and R. consanguinea var. rigida and the closely related R. emaciata, R. riparia and R. tenuis from section Tenues. On the other hand, karyotypic similarities occurred among sections Valderugosae, Marisculae, Stenophylku and Cemuae.     

Chromosome numbers and karyotypes of South American species and populations of Hypochaeris (Asteraceae)

One hundred and thirty-seven new chromosome counts are reported from 104 populations of 26 native South American taxa of Hypochaeris (Asteraceae, Lactuceae), together with two invasive Mediterranean species: H. glabra and H. radicata . First reports are provided for seven taxa ( H. alba , H. cf. eremophila , H. caespitosa , H. hookeri , H. parodii , H. patagonica and H. pinnatifida ) and one new ploidy level is reported (diploid for H. incana , so far known only as a tetraploid). Including the results of this study, the chromosomes of 39 of the c . 50 Hypochaeris species known from the New World have now been counted. Most species are diploid with 2 n  = 2 x  = 8 and have bimodal, asymmetrical karyotypes. Tetraploidy (2 n  = 4 x  = 16) is reported here for the first time in H. caespitosa . Infra-specific polyploidy (probably autopolyploidy) is reported in H. incana and H. taraxacoides , both cases including infra-populational cytotype mixtures (2 x and 4 x ). Polyploidy is now known from eight South American Hypochaeris species ( c . 16%). Basic karyotype analyses allow the placement of the newly counted taxa into previously proposed but slightly modified groupings and provide the framework for further molecular cytogenetic analyses. The reported findings suggest that chromosomal change in South American Hypochaeris , in contrast to Old World species, has not involved aneuploidy, but polyploidy and/or more subtle changes in chromosome length, perhaps via satellite DNA amplification/deletion or activity of retroelements, and rDNA reorganization.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 49–60.     

Chromosome numbers in Hieracium and Pilosella species (Asteraceae) from the Iberian Peninsula and the Balearic Islands

Mitotic chromosome numbers are reported from 16 Hieracium and two Pilosella species from the Iberian Peninsula and the Balearic Islands that are little known cytogenetically. The chromosome numbers of Hieracium aragonense Scheele (2 n  = 27, 36), H. compositum Lapeyr. (2 n  = 27), H. murcandidum G. Mateo (2 n  = 27), H. spathulatum Scheele (2 n  = 27), H. segurae Mateo (2 n  = 27), H. teruelanum Mateo (2 n  = 27), H. valentinum Pau (2 n  = 27), Pilosella pseudovahlii (De Retz) Mateo (2 n  = 18), and P. tardans (Peter) Sójak (2 n  = 36) were determined for the first time. New cytotypes were detected in H. cordifolium Lapeyr. (2 n  = 27) and H. loscosianum Scheele (2 n  = 36). The karyotype of the studied species was similar in overall morphology and comprised metacentric and submetacentric chromosomes; secondary constrictions (up to two) were detected in the chromosome complements of some species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 311–320.     

Gradual genome size evolution and polyploidy in Allium from the Qinghai–Tibetan Plateau

Background and AimsGenome size is an important plant trait, with substantial interspecies variation. The mechanisms and selective pressures underlying genome size evolution are important topics in evolutionary biology. There is considerable diversity in Allium from the Qinghai–Tibetan Plateau, where genome size variation and related evolutionary mechanisms are poorly understood.MethodsWe reconstructed the Allium phylogeny using DNA sequences from 71 species. We also estimated genome sizes of 62 species, and determined chromosome numbers in 65 species. We examined the phylogenetic signal associated with genome size variation, and tested how well the data fit different evolutionary models. Correlations between genome size variations and seed mass, altitude and 19 bioclimatic factors were determined.Key Results Allium genome sizes differed substantially between species and within diploids, triploids, tetraploids, hexaploids and octaploids. Size per monoploid genome (1Cx) tended to decrease with increasing ploidy levels. Allium polyploids tended to grow at a higher altitude than diploids. The phylogenetic tree was divided into three evolutionary branches. The genomes in Clade I were mostly close to the ancestral genome (18.781 pg) while those in Clades II and III tended to expand and contract, respectively. A weak phylogenetic signal was detected for Allium genome size. Furthermore, significant positive correlations were detected between genome size and seed mass, as well as between genome size and altitude. However, genome size was not correlated with 19 bioclimatic variables.Conclusions Allium genome size shows gradual evolution, followed by subsequent adaptive radiation. The three well-supported Allium clades are consistent with previous studies. The evolutionary patterns in different Allium clades revealed genome contraction, expansion and relative stasis. The Allium species in Clade II may follow adaptive radiation. The genome contraction in Clade III may be due to DNA loss after polyploidization. Allium genome size might be influenced by selective pressure due to the conditions on the Qinghai–Tibetan Plateau (low temperature, high UV irradiation and abundant phosphate in the soil).     

Occurrence of tetraploid and octoploid cytotypes in Senecio jacobaea ssp. jacobaea (Asteraceae) in Pannonia and the Carpathians

Four different cytotypes have been reported for Senecio jacobaea L. ssp. jacobaea throughout Europe, with the most common occurrence of tetraploids (2 n  = 40). Here we present a survey of previously published chromosome number data on this subspecies and its geographical distribution, and focus on populations from Pannonia and the Carpathians. Two ploidy levels have been determined in the study area, using chromosome counting and flow cytometry: tetraploid (2 n  = 40) and octoploid (2 n  = 80). Fifty-one populations originating from Slovakia, Austria, the Czech Republic, Hungary, Ukraine and Romania have been analysed. Multivariate morphometric analyses have been performed on 39 populations to study morphological differentiation between these two cytotypes. Despite slight morphological tendencies expressed on the level of populations, tetraploid and octoploids cannot be reliably distinguished morphologically and they are not taxonomically classified formally here.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 231–242.     

New or rarely reported chromosome numbers in taxa of subtribe Artemisiinae (Anthemideae, Asteraceae) from Mongolia

This study encompasses 25 chromosome counts of 18 species in the subtribe Artemisiinae (tribe Anthemideae) of the family Asteraceae, from Mongolia. Most (15 species) belong to Artemisia , the largest genus of the subtribe, whereas the others come from two genera very closely related to it: Ajania (two species) and Neopallasia (one species). Eleven counts are new reports, three are not consistent with previous reports and the remainder confirm scanty earlier information. The majority of the species have x  = 9 as their basic chromosome number, but there are some taxa with x  = 8. Ploidy levels range from 2 x to 6 x . The presence of B-chromosomes was detected in Ajania fruticulosa .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 203–210.