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Papers by barbora singliarova

TAXON

Members of the Pilosella alpicola species group (Asteraceae) are distributed throughout the alpin... more Members of the Pilosella alpicola species group (Asteraceae) are distributed throughout the alpine region of the European mountains (Alps, Carpathians, Balkan mountains). Like other Pilosella species groups (Hieracium subg. Pilosella), the taxonomy and species' relationships are poorly understood mostly due to widespread facultative apomixis, frequent hybridization and polyploidization-the most important phenomena substantially involved in the evolutionary history of the genus. We assessed morphology, ploidy level, variation in breeding system and molecular variation within the P. alpicola group to provide a new taxonomic concept and to clarify evolutionary relationships among species and origin of polyploids. Multivariate morphometric analyses (UPGMA, CDA, PCA) applied on 324 plants originated from 21 populations revealed existence of four well-separated clusters corresponding to four allopatric taxa: P. alpicola s.str. (Alps), P. rhodopea (Balkan Peninsula, Southern Carpathians), P. serbica (Serbia and Montenegro) and P. ullepitschii (Carpathians). In total, four ploidy levels (2x, 3x, 4x, 5x) were detected among 557 plants analysed from 19 populations by classical chromosome counting and flow cytometric analysis. The tetra-and pentaploids of P. alpicola s.str. have an allopatric distribution (Wallis Alps vs. Dolomites, respectively). Four ploidy levels with complex cytogeographic pattern and high frequency of mixed ploidy populations (75%) indicating a primary contact zone were recorded in P. rhodopea. Pilosella ullepitschii and P. serbica are exclusively diploid and are both sexually reproducing. In spite of a clear morphological separation, a molecular analysis (ITS and cpDNA sequences) suggests close relationships and rather recent origin of all studied taxa, except P. alpicola s.str. The latter taxon is an agamospermic allopolyploid that likely originated polytopically from a hybridization between P. rhodopea from the Balkans and P. glacialis from the Alps. In contrast to P. alpicola s.str., our data strongly support an autopolyploid origin of P. rhodopea polyploids which reproduced strictly sexually. Pilosella petraea, sometimes treated as a member of the P. alpicola group, differs from the remaining taxa by its conspicuous morphology, ecology and ITS polymorphism and should be removed from the group. Range shifts and extinctions were likely involved in shaping the evolutionary and modern distributional pattern of the group. Our combined methodological approach enabled us to propose a new taxonomic circumscription for the P. alpicola group and revealed auto-and allopolyploidization events.

Annals of Botany, Apr 13, 2012

† Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead t... more † Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead to the elimination of the less common cytotype, unless prezygotic barriers enhance assortative mating. However, such barriers favouring cytotype coexistence have only rarely been explored. Here, an assessment is made of the mechanisms involved in formation of mixed-ploidy populations and coexistence of diploid plants and their closely related allotetraploid derivates from the Centaurea stoebe complex (Asteraceae). † Methods An investigation was made of microspatial and microhabitat distribution, life-history and fitness traits, flowering phenology, genetic relatedness of cytotypes and intercytotype gene flow (cpDNA and microsatellites) in six mixed-ploidy populations in Central Europe. † Key Results Diploids and tetraploids were genetically differentiated, thus corroborating the secondary origin of contact zones. The cytotypes were spatially segregated at all sites studied, with tetraploids colonizing preferentially drier and open microhabitats created by human-induced disturbances. Conversely, they were rare in more natural microsites and microsites with denser vegetation despite their superior persistence ability (polycarpic life cycle). The seed set of tetraploid plants was strongly influenced by their frequency in mixed-ploidy populations. Triploid hybrids originated from bidirectional hybridizations were extremely rare and almost completely sterile, indicating a strong postzygotic barrier between cytotypes. † Conclusions The findings suggest that tetraploids are later immigrants into already established diploid populations and that anthropogenic activities creating open niches favouring propagule introductions were the major factor shaping the non-random distribution and habitat segregation of cytotypes at fine spatial scale. Establishment and spread of tetraploids was further facilitated by their superior persistence through the perennial life cycle. The results highlight the importance of non-adaptive spatio-temporal processes in explaining microhabitat and microspatial segregation of cytotypes.

Nordic Journal of Botany, Feb 1, 2013

ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically ... more ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically vicariant alpine taxa. We performed a thorough herbarium revision and literature survey to infer their distributional pattern(s). Pilosella alpicola s.s. occurs in the Alps in two disjunct areas: the Swiss Valais Alps and the Italian Dolomites. Historical records come also from the Austrian Alps (Gurktaler Alps and Hohe Tauern) and from one site from the Alpes Maritimes (Col de Larche), but the localities have not been recently confirmed. Pilosella rhodopea, a Balkan subendemic taxon, is quite widespread in Bulgaria (Stara planina Mts, Rila Mts and Pirin Mts), but is more rare in Albania, Greece and Macedonia. Interestingly, this species has also been recorded at two isolated sites in the Romanian southern Carpathians (the Căpăţînii and Cozia Mts). This occurrence underlines the floristic affinities of this part of the Carpathians to the Balkan flora. Only two localities of P. serbica, based on voucher specimens, have been recorded so far; Kopaonik Mts in Serbia and the Prokletije Mts in Montenegro. The records from other ranges are related to P. rhodopea. Pilosella ullepitschii, the detailed distribution of which has already been published, is a Carpathian endemic with its core area of distribution in the western Carpathians (Slovakia and Poland). Three isolated localities are also known in the eastern Carpathians (Nemira Mts) and one locality in the southern Carpathians (Bucegi Mts). The possible causes of disjunctions between and within species ranges are briefly discussed. Based on the distributional data, population sizes and ecology, we evaluate the conservation status of the P. alpicola taxa and propose their inclusion in national Red Lists.

Applications in Plant Sciences, 2015

Microsatellite markers were developed for the Pilosella alpicola group (Asteraceae), comprising f... more Microsatellite markers were developed for the Pilosella alpicola group (Asteraceae), comprising four closely related species distributed in subalpine areas of Europe. These species are believed to have diverged recently, but display contrasting cytogeographic patterns and variation in breeding systems, representing a promising model system for studying plant speciation, adaptation, and recent polyploidization. We developed 17 microsatellite markers for the P. alpicola group using 454 sequencing. Sixteen markers were polymorphic, with the number of alleles per locus ranging from seven to 16 and observed and expected heterozygosity ranging from 0.45 to 0.84 and 0.72 to 0.92, respectively. Ten and five loci amplified in the related species, P. echioides and P. officinarum, respectively, but only two in Andryala and one in Hieracium s. str. The developed microsatellite markers have high potential to become useful tools to study microevolutionary processes in the P. alpicola group and re...

Preslia -Praha-

Exploring patterns of variation within the central-European Tephroseris longifolia agg.: karyolog... more Exploring patterns of variation within the central-European Tephroseris longifolia agg.: karyological and morphological study Karyologická a morfologická variabilita v rámci Tephroseris longifolia agg.

Applications in Plant Sciences, 2015

BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access t... more BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses.

Nordic Journal of Botany, 2007

Chromosome numbers for 88 plants of Picris hieracioides s.l. originating from 32 localities from ... more Chromosome numbers for 88 plants of Picris hieracioides s.l. originating from 32 localities from Austria, Andorra, Croatia, France, Germany, Hungary, Italy, Slovakia and Spain are reported. All analysed plants were diploid, with 2n" 2x" 10. In addition to P. hieracioides s.l., we also found diploidy (2n" 2x" 10) in P. nuristanica from Kirgizia and P. japonica from Japan. Our data

, PL = ploidy level detected by flow cytometry. Chromosome numbers of the plants marked by '!' ha... more , PL = ploidy level detected by flow cytometry. Chromosome numbers of the plants marked by '!' have been previously published in Rotreklová et al.

Botanica Helvetica

Hieracium alpinum L. (Asteraceae) is an arctic-alpine species distributed throughout Europe with ... more Hieracium alpinum L. (Asteraceae) is an arctic-alpine species distributed throughout Europe with both diploid and triploid cytotypes. We determined the ploidy levels of plants from 23 populations from Austria, Bosnia and Herzegovina, Finland, Italy, Norway, Romania, Slovakia, Switzerland and Ukraine. Data showed a non-overlapping pattern of cytotype distribution: sexually reproducing diploids (2n = 2x = 18) occur solely in the Eastern and Southern Carpathians, while apomictic triploids (2n = 3x = 27) cover the rest of the range. Such clear-cut allopatry is rather rare in vascular plants with geographical parthenogenesis. Comparison of absolute genome size indicates genome downsizing (by on average 3.7%) of haploid DNA amount in triploids relative to diploids. Genome size further correlated with longitude and latitude in the Alps, with decreasing absolute DNA content from west to east, and from south to north. While previously published data indicated complete male sterility of tripl...

Annals of botany, 2008

Pilosella officinarum (syn. Hieracium pilosella) is a highly structured species with respect to t... more Pilosella officinarum (syn. Hieracium pilosella) is a highly structured species with respect to the ploidy level, with obvious cytogeographic trends. Previous non-collated data indicated a possible differentiation in the frequency of particular ploidy levels in the Czech Republic and Slovakia. Therefore, detailed sampling and ploidy level analyses were assessed to reveal a boundary of common occurrence of tetraploids on one hand and higher ploids on the other. For a better understanding of cytogeographic differentiation of P. officinarum in central Europe, a search was made for a general cytogeographic pattern in Europe based on published data. DNA-ploidy level and/or chromosome number were identified for 1059 plants using flow cytometry and/or chromosome counting on root meristem preparations. Samples were collected from 336 localities in the Czech Republic, Slovakia and north-eastern Hungary. In addition, ploidy levels were determined for plants from 18 localities in Bulgaria, Geo...

Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian ... more Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian endemic. Its distribution range comprises two areas separated by about 600 km. While in the Western Carpathians (Slovakia and Poland) the taxon occurs in numerous sites, only four localities of man-made origin are known from the Eastern and Southern Carpathians (Romania). We used allozyme markers to test two likely possible scenarios for the origin of this disjunction: long distance dispersal and vicariance. Our data indicate a significant loss of genetic diversity in the isolated Eastern and Southern Carpathian populations in following genetic parameters (averaged per region): percentage of polymorphic loci (38.9% found in the Eastern and Southern Carpathians versus 58.3% in the Western Carpathians), allelic richness (1.4 vs. 1.6), expected heterozygosity (0.134 vs. 0.235), mean number of distinguishable multilocus genotypes (4.3 vs. 10.6) and proportion of distinguishable multilocus genotypes (0.34 vs. 0.68). Higher proportion of homozygous loci found in the Eastern and Southern Carpathian populations might indicate a higher rate of inbreeding due to non-random mating. We assume that these genetically depauperate populations have experienced a very strong genetic bottleneck, probably due to a founder effect. Although our data suggest that the longdistance dispersal model is most likely, more discriminate genetic markers should be used to test this further.

Annals of Botany, 2012

† Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead t... more † Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead to the elimination of the less common cytotype, unless prezygotic barriers enhance assortative mating. However, such barriers favouring cytotype coexistence have only rarely been explored. Here, an assessment is made of the mechanisms involved in formation of mixed-ploidy populations and coexistence of diploid plants and their closely related allotetraploid derivates from the Centaurea stoebe complex (Asteraceae). † Methods An investigation was made of microspatial and microhabitat distribution, life-history and fitness traits, flowering phenology, genetic relatedness of cytotypes and intercytotype gene flow (cpDNA and microsatellites) in six mixed-ploidy populations in Central Europe. † Key Results Diploids and tetraploids were genetically differentiated, thus corroborating the secondary origin of contact zones. The cytotypes were spatially segregated at all sites studied, with tetraploids colonizing preferentially drier and open microhabitats created by human-induced disturbances. Conversely, they were rare in more natural microsites and microsites with denser vegetation despite their superior persistence ability (polycarpic life cycle). The seed set of tetraploid plants was strongly influenced by their frequency in mixed-ploidy populations. Triploid hybrids originated from bidirectional hybridizations were extremely rare and almost completely sterile, indicating a strong postzygotic barrier between cytotypes. † Conclusions The findings suggest that tetraploids are later immigrants into already established diploid populations and that anthropogenic activities creating open niches favouring propagule introductions were the major factor shaping the non-random distribution and habitat segregation of cytotypes at fine spatial scale. Establishment and spread of tetraploids was further facilitated by their superior persistence through the perennial life cycle. The results highlight the importance of non-adaptive spatio-temporal processes in explaining microhabitat and microspatial segregation of cytotypes.

A taxonomic study of the Pilosella alpicola group growing in the Carpathians revealed the presenc... more A taxonomic study of the Pilosella alpicola group growing in the Carpathians revealed the presence of two morphologically distinguishable taxa: P. ullepitschii (Błocki) Szeląg and P. rhodopea (Griseb.) Szeląg. While P. ullepitschii is endemic to the Carpathians, P. rhodopea is a Balkan subendemic with two isolated localities in the Southern Carpathians (Mt Cozia and Mt Zmeuretu). The core area of distribution of P. ullepitchii is the natural subalpine and alpine meadows of the Western Carpathians (the Vysoké and Západné Tatry Mts in Slovakia and Poland). In addition, only three isolated localities are known from the Nemira Mts (Romanian Eastern Carpathians) and one from the Bucegi Mts (Romanian Southern Carpathians). Interestingly, the Romanian populations occur in man-made habitats (secondary pastures). Karyological and flow cytometric analyses of 305 plants from 13 populations of P. ullepitschii revealed only diploid plants (2n = 2x = 18). One Carpathian population of P. rhodopea from Mt Cozia is also diploid. This is the first report of diploidy in this species. However, the populations from the main part of the distribution of this taxon in the Balkan mountains include other cytotypes. Detailed morphological descriptions and distributions for both taxa are given.

Background and Aims Pilosella officinarum (syn. Hieracium pilosella) is a highly structured speci... more Background and Aims Pilosella officinarum (syn. Hieracium pilosella) is a highly structured species with respect to the ploidy level, with obvious cytogeographic trends. Previous non-collated data indicated a possible differentiation in the frequency of particular ploidy levels in the Czech Republic and Slovakia. Therefore, detailed sampling and ploidy level analyses were assessed to reveal a boundary of common occurrence of tetraploids on one hand and higher ploids on the other. For a better understanding of cytogeographic differentiation of P. officinarum in central Europe, a search was made for a general cytogeographic pattern in Europe based on published data. † Methods DNA-ploidy level and/or chromosome number were identified for 1059 plants using flow cytometry and/or chromosome counting on root meristem preparations. Samples were collected from 336 localities in the Czech Republic, Slovakia and northeastern Hungary. In addition, ploidy levels were determined for plants from 18 localities in Bulgaria, Georgia, Ireland, Italy, Romania and Ukraine. † Key Results Four ploidy levels were found in the studied area with a contrasting pattern of distribution. The most widespread cytotype in the western part of the Czech Republic is tetraploid (4x) reproducing sexually, while the apomictic pentaploids and mostly apomictic hexaploids (5x and 6x, respectively) clearly prevail in Slovakia and the eastern part of the Czech Republic. The boundary between common occurrence of tetraploids and higher ploids is very obvious and represents the geomorphologic boundary between the Bohemian Massif and the Western Carpathians with the adjacent part of Pannonia. Mixed populations consisting of two different ploidy levels were recorded in nearly 11% of localities. A statistically significant difference in a vertical distribution of penta-and hexaploids was observed in the Western Carpathians and the adjacent Pannonian Plain. Hexaploid populations tend to occur at lower elevations (usually below 500 m), while the pentaploid level is more or less evenly distributed up to 1000 m a.s.l. For the first time the heptaploid level (7x) was found on one site in Slovakia. In Europe, the sexual tetraploid level has clearly a sub-Atlantic character of distribution. The plants of higher ploidy level (penta-and hexa-) with mostly apomictic reproduction prevail in the northern part of Scandinavia and the British Isles, the Alps and the Western Carpathians with the adjacent part of Pannonia. A detailed overview of published data shows that extremely rare records on existence of diploid populations in the southwest Alps are with high probability erroneous and most probably refer to the closely related diploid species P. peleteriana. † Conclusions The recent distribution of P. officinarum in Europe is complex and probably reflects the climatic changes during the Pleistocene and consequent postglacial migrations. Probably both penta-and hexaploids arose independently in central Europe (Alps and Carpathian Mountains) and in northern Europe (Scandinavia, Great Britain, Ireland), where the apomictic plants colonized deglaciated areas. We suggest that P. officinarum is in fact an amphidiploid species with a basic tetraploid level, which probably originated from hybridizations of diploid taxa from the section Pilosellina.

Plant Systematics and Evolution, 2012

The genome size of 265 plants and the GC content of 126 plants from 63 populations of the Cyanus ... more The genome size of 265 plants and the GC content of 126 plants from 63 populations of the Cyanus triumfetti and Cyanus montanus groups, collected across the Carpathians, Pannonia, Bohemian Massif, and Western and Dinaric Alps were determined by PI and DAPI flow cytometry. Variation of the nuclear DNA content among homoploid species, and intraspecific and interpopulation variation were confirmed in simultaneous analyses. The 2C-value at the diploid level (the C. triumfetti group) varied from 2.53 for Cyanus dominii subsp. sokolensis to 3.06 pg for C. triumfetti s.s. (1.21-fold range). At the tetraploid level (the C. montanus group), the 2C-value varied from 5.19 for Cyanus mollis to 5.84 pg for C. montanus (1.13-fold range). High intraspecific and interpopulation variation in the amount of nuclear DNA in the C. triumfetti group correlates with the extensive morphological variation found in this group. Significant between-species differences in genome size indicate that this attribute may be used as a supportive taxonomic marker for both of the groups studied. The GC content varied by 2.93 %, from 39.46 % for ''Cyanus axillaris'' to 40.61 % for Cyanus adscendens; this character is of no value for taxonomic purposes. Genome size of the studied populations is significantly higher in southern parts of the distribution area and at higher elevations. Plants with smaller genomes tend to occur in dry areas at low altitudes with high diurnal and annual temperature oscillations. The GC content of the populations studied is significantly correlated with longitude, increasing from east to west; and plants with GC-rich genomes are concentrated in the coldest areas with low minimum temperatures. Keywords Base composition Á Cyanus triumfetti group Á Environmental conditions Á Flow cytometry Á Homoploid species Á Taxonomy Electronic supplementary material The online version of this article (

Plant Systematics and Evolution, 2008

Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian ... more Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian endemic. Its distribution range comprises two areas separated by about 600 km. While in the Western Carpathians (Slovakia and Poland) the taxon occurs in numerous sites, only four localities of man-made origin are known from the Eastern and Southern Carpathians (Romania). We used allozyme markers to test two likely possible scenarios for the origin of this disjunction: long distance dispersal and vicariance. Our data indicate a significant loss of genetic diversity in the isolated Eastern and Southern Carpathian populations in following genetic parameters (averaged per region): percentage of polymorphic loci (38.9% found in the Eastern and Southern Carpathians versus 58.3% in the Western Carpathians), allelic richness (1.4 vs. 1.6), expected heterozygosity (0.134 vs. 0.235), mean number of distinguishable multilocus genotypes (4.3 vs. 10.6) and proportion of distinguishable multilocus genotypes (0.34 vs. 0.68). Higher proportion of homozygous loci found in the Eastern and Southern Carpathian populations might indicate a higher rate of inbreeding due to non-random mating. We assume that these genetically depauperate populations have experienced a very strong genetic bottleneck, probably due to a founder effect. Although our data suggest that the longdistance dispersal model is most likely, more discriminate genetic markers should be used to test this further.

Nordic Journal of Botany, 2007

and Spain are reported. All analysed plants were diploid, with 2n 0 2x010. In addition to P. hier... more and Spain are reported. All analysed plants were diploid, with 2n 0 2x010. In addition to P. hieracioides s.l., we also found diploidy (2n 02x010) in P. nuristanica from Kirgizia and P. japonica from Japan. Our data confirmed previously published results in the literature. We also studied the mode of reproduction in all three taxa. Picris nuristanica and P. japonica s.l. were able to produce some seeds by selfing, and this is the first record of autogamy in this genus. In P. hieracioides s.l. only strict allogamy was found, demonstrating the presence of a self-incompatibility system in this species. Because apomixis, namely apospory, has previously been reported in P. hieracioides, castration experiments were performed in order to confirm or reject this report. However, no evidence of apomictic seed formation was found within this species. In the light of our results, P. hieracioides should be considered a strictly sexual taxon.

Nordic Journal of Botany, 2013

ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically ... more ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically vicariant alpine taxa. We performed a thorough herbarium revision and literature survey to infer their distributional pattern(s). Pilosella alpicola s.s. occurs in the Alps in two disjunct areas: the Swiss Valais Alps and the Italian Dolomites. Historical records come also from the Austrian Alps (Gurktaler Alps and Hohe Tauern) and from one site from the Alpes Maritimes (Col de Larche), but the localities have not been recently confirmed. Pilosella rhodopea, a Balkan subendemic taxon, is quite widespread in Bulgaria (Stara planina Mts, Rila Mts and Pirin Mts), but is more rare in Albania, Greece and Macedonia. Interestingly, this species has also been recorded at two isolated sites in the Romanian southern Carpathians (the Căpăţînii and Cozia Mts). This occurrence underlines the floristic affinities of this part of the Carpathians to the Balkan flora. Only two localities of P. serbica, based on voucher specimens, have been recorded so far; Kopaonik Mts in Serbia and the Prokletije Mts in Montenegro. The records from other ranges are related to P. rhodopea. Pilosella ullepitschii, the detailed distribution of which has already been published, is a Carpathian endemic with its core area of distribution in the western Carpathians (Slovakia and Poland). Three isolated localities are also known in the eastern Carpathians (Nemira Mts) and one locality in the southern Carpathians (Bucegi Mts). The possible causes of disjunctions between and within species ranges are briefly discussed. Based on the distributional data, population sizes and ecology, we evaluate the conservation status of the P. alpicola taxa and propose their inclusion in national Red Lists.

TAXON

Members of the Pilosella alpicola species group (Asteraceae) are distributed throughout the alpin... more Members of the Pilosella alpicola species group (Asteraceae) are distributed throughout the alpine region of the European mountains (Alps, Carpathians, Balkan mountains). Like other Pilosella species groups (Hieracium subg. Pilosella), the taxonomy and species' relationships are poorly understood mostly due to widespread facultative apomixis, frequent hybridization and polyploidization-the most important phenomena substantially involved in the evolutionary history of the genus. We assessed morphology, ploidy level, variation in breeding system and molecular variation within the P. alpicola group to provide a new taxonomic concept and to clarify evolutionary relationships among species and origin of polyploids. Multivariate morphometric analyses (UPGMA, CDA, PCA) applied on 324 plants originated from 21 populations revealed existence of four well-separated clusters corresponding to four allopatric taxa: P. alpicola s.str. (Alps), P. rhodopea (Balkan Peninsula, Southern Carpathians), P. serbica (Serbia and Montenegro) and P. ullepitschii (Carpathians). In total, four ploidy levels (2x, 3x, 4x, 5x) were detected among 557 plants analysed from 19 populations by classical chromosome counting and flow cytometric analysis. The tetra-and pentaploids of P. alpicola s.str. have an allopatric distribution (Wallis Alps vs. Dolomites, respectively). Four ploidy levels with complex cytogeographic pattern and high frequency of mixed ploidy populations (75%) indicating a primary contact zone were recorded in P. rhodopea. Pilosella ullepitschii and P. serbica are exclusively diploid and are both sexually reproducing. In spite of a clear morphological separation, a molecular analysis (ITS and cpDNA sequences) suggests close relationships and rather recent origin of all studied taxa, except P. alpicola s.str. The latter taxon is an agamospermic allopolyploid that likely originated polytopically from a hybridization between P. rhodopea from the Balkans and P. glacialis from the Alps. In contrast to P. alpicola s.str., our data strongly support an autopolyploid origin of P. rhodopea polyploids which reproduced strictly sexually. Pilosella petraea, sometimes treated as a member of the P. alpicola group, differs from the remaining taxa by its conspicuous morphology, ecology and ITS polymorphism and should be removed from the group. Range shifts and extinctions were likely involved in shaping the evolutionary and modern distributional pattern of the group. Our combined methodological approach enabled us to propose a new taxonomic circumscription for the P. alpicola group and revealed auto-and allopolyploidization events.

Annals of Botany, Apr 13, 2012

† Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead t... more † Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead to the elimination of the less common cytotype, unless prezygotic barriers enhance assortative mating. However, such barriers favouring cytotype coexistence have only rarely been explored. Here, an assessment is made of the mechanisms involved in formation of mixed-ploidy populations and coexistence of diploid plants and their closely related allotetraploid derivates from the Centaurea stoebe complex (Asteraceae). † Methods An investigation was made of microspatial and microhabitat distribution, life-history and fitness traits, flowering phenology, genetic relatedness of cytotypes and intercytotype gene flow (cpDNA and microsatellites) in six mixed-ploidy populations in Central Europe. † Key Results Diploids and tetraploids were genetically differentiated, thus corroborating the secondary origin of contact zones. The cytotypes were spatially segregated at all sites studied, with tetraploids colonizing preferentially drier and open microhabitats created by human-induced disturbances. Conversely, they were rare in more natural microsites and microsites with denser vegetation despite their superior persistence ability (polycarpic life cycle). The seed set of tetraploid plants was strongly influenced by their frequency in mixed-ploidy populations. Triploid hybrids originated from bidirectional hybridizations were extremely rare and almost completely sterile, indicating a strong postzygotic barrier between cytotypes. † Conclusions The findings suggest that tetraploids are later immigrants into already established diploid populations and that anthropogenic activities creating open niches favouring propagule introductions were the major factor shaping the non-random distribution and habitat segregation of cytotypes at fine spatial scale. Establishment and spread of tetraploids was further facilitated by their superior persistence through the perennial life cycle. The results highlight the importance of non-adaptive spatio-temporal processes in explaining microhabitat and microspatial segregation of cytotypes.

Nordic Journal of Botany, Feb 1, 2013

ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically ... more ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically vicariant alpine taxa. We performed a thorough herbarium revision and literature survey to infer their distributional pattern(s). Pilosella alpicola s.s. occurs in the Alps in two disjunct areas: the Swiss Valais Alps and the Italian Dolomites. Historical records come also from the Austrian Alps (Gurktaler Alps and Hohe Tauern) and from one site from the Alpes Maritimes (Col de Larche), but the localities have not been recently confirmed. Pilosella rhodopea, a Balkan subendemic taxon, is quite widespread in Bulgaria (Stara planina Mts, Rila Mts and Pirin Mts), but is more rare in Albania, Greece and Macedonia. Interestingly, this species has also been recorded at two isolated sites in the Romanian southern Carpathians (the Căpăţînii and Cozia Mts). This occurrence underlines the floristic affinities of this part of the Carpathians to the Balkan flora. Only two localities of P. serbica, based on voucher specimens, have been recorded so far; Kopaonik Mts in Serbia and the Prokletije Mts in Montenegro. The records from other ranges are related to P. rhodopea. Pilosella ullepitschii, the detailed distribution of which has already been published, is a Carpathian endemic with its core area of distribution in the western Carpathians (Slovakia and Poland). Three isolated localities are also known in the eastern Carpathians (Nemira Mts) and one locality in the southern Carpathians (Bucegi Mts). The possible causes of disjunctions between and within species ranges are briefly discussed. Based on the distributional data, population sizes and ecology, we evaluate the conservation status of the P. alpicola taxa and propose their inclusion in national Red Lists.

Applications in Plant Sciences, 2015

Microsatellite markers were developed for the Pilosella alpicola group (Asteraceae), comprising f... more Microsatellite markers were developed for the Pilosella alpicola group (Asteraceae), comprising four closely related species distributed in subalpine areas of Europe. These species are believed to have diverged recently, but display contrasting cytogeographic patterns and variation in breeding systems, representing a promising model system for studying plant speciation, adaptation, and recent polyploidization. We developed 17 microsatellite markers for the P. alpicola group using 454 sequencing. Sixteen markers were polymorphic, with the number of alleles per locus ranging from seven to 16 and observed and expected heterozygosity ranging from 0.45 to 0.84 and 0.72 to 0.92, respectively. Ten and five loci amplified in the related species, P. echioides and P. officinarum, respectively, but only two in Andryala and one in Hieracium s. str. The developed microsatellite markers have high potential to become useful tools to study microevolutionary processes in the P. alpicola group and re...

Preslia -Praha-

Exploring patterns of variation within the central-European Tephroseris longifolia agg.: karyolog... more Exploring patterns of variation within the central-European Tephroseris longifolia agg.: karyological and morphological study Karyologická a morfologická variabilita v rámci Tephroseris longifolia agg.

Applications in Plant Sciences, 2015

BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access t... more BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses.

Nordic Journal of Botany, 2007

Chromosome numbers for 88 plants of Picris hieracioides s.l. originating from 32 localities from ... more Chromosome numbers for 88 plants of Picris hieracioides s.l. originating from 32 localities from Austria, Andorra, Croatia, France, Germany, Hungary, Italy, Slovakia and Spain are reported. All analysed plants were diploid, with 2n" 2x" 10. In addition to P. hieracioides s.l., we also found diploidy (2n" 2x" 10) in P. nuristanica from Kirgizia and P. japonica from Japan. Our data

, PL = ploidy level detected by flow cytometry. Chromosome numbers of the plants marked by '!' ha... more , PL = ploidy level detected by flow cytometry. Chromosome numbers of the plants marked by '!' have been previously published in Rotreklová et al.

Botanica Helvetica

Hieracium alpinum L. (Asteraceae) is an arctic-alpine species distributed throughout Europe with ... more Hieracium alpinum L. (Asteraceae) is an arctic-alpine species distributed throughout Europe with both diploid and triploid cytotypes. We determined the ploidy levels of plants from 23 populations from Austria, Bosnia and Herzegovina, Finland, Italy, Norway, Romania, Slovakia, Switzerland and Ukraine. Data showed a non-overlapping pattern of cytotype distribution: sexually reproducing diploids (2n = 2x = 18) occur solely in the Eastern and Southern Carpathians, while apomictic triploids (2n = 3x = 27) cover the rest of the range. Such clear-cut allopatry is rather rare in vascular plants with geographical parthenogenesis. Comparison of absolute genome size indicates genome downsizing (by on average 3.7%) of haploid DNA amount in triploids relative to diploids. Genome size further correlated with longitude and latitude in the Alps, with decreasing absolute DNA content from west to east, and from south to north. While previously published data indicated complete male sterility of tripl...

Annals of botany, 2008

Pilosella officinarum (syn. Hieracium pilosella) is a highly structured species with respect to t... more Pilosella officinarum (syn. Hieracium pilosella) is a highly structured species with respect to the ploidy level, with obvious cytogeographic trends. Previous non-collated data indicated a possible differentiation in the frequency of particular ploidy levels in the Czech Republic and Slovakia. Therefore, detailed sampling and ploidy level analyses were assessed to reveal a boundary of common occurrence of tetraploids on one hand and higher ploids on the other. For a better understanding of cytogeographic differentiation of P. officinarum in central Europe, a search was made for a general cytogeographic pattern in Europe based on published data. DNA-ploidy level and/or chromosome number were identified for 1059 plants using flow cytometry and/or chromosome counting on root meristem preparations. Samples were collected from 336 localities in the Czech Republic, Slovakia and north-eastern Hungary. In addition, ploidy levels were determined for plants from 18 localities in Bulgaria, Geo...

Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian ... more Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian endemic. Its distribution range comprises two areas separated by about 600 km. While in the Western Carpathians (Slovakia and Poland) the taxon occurs in numerous sites, only four localities of man-made origin are known from the Eastern and Southern Carpathians (Romania). We used allozyme markers to test two likely possible scenarios for the origin of this disjunction: long distance dispersal and vicariance. Our data indicate a significant loss of genetic diversity in the isolated Eastern and Southern Carpathian populations in following genetic parameters (averaged per region): percentage of polymorphic loci (38.9% found in the Eastern and Southern Carpathians versus 58.3% in the Western Carpathians), allelic richness (1.4 vs. 1.6), expected heterozygosity (0.134 vs. 0.235), mean number of distinguishable multilocus genotypes (4.3 vs. 10.6) and proportion of distinguishable multilocus genotypes (0.34 vs. 0.68). Higher proportion of homozygous loci found in the Eastern and Southern Carpathian populations might indicate a higher rate of inbreeding due to non-random mating. We assume that these genetically depauperate populations have experienced a very strong genetic bottleneck, probably due to a founder effect. Although our data suggest that the longdistance dispersal model is most likely, more discriminate genetic markers should be used to test this further.

Annals of Botany, 2012

† Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead t... more † Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead to the elimination of the less common cytotype, unless prezygotic barriers enhance assortative mating. However, such barriers favouring cytotype coexistence have only rarely been explored. Here, an assessment is made of the mechanisms involved in formation of mixed-ploidy populations and coexistence of diploid plants and their closely related allotetraploid derivates from the Centaurea stoebe complex (Asteraceae). † Methods An investigation was made of microspatial and microhabitat distribution, life-history and fitness traits, flowering phenology, genetic relatedness of cytotypes and intercytotype gene flow (cpDNA and microsatellites) in six mixed-ploidy populations in Central Europe. † Key Results Diploids and tetraploids were genetically differentiated, thus corroborating the secondary origin of contact zones. The cytotypes were spatially segregated at all sites studied, with tetraploids colonizing preferentially drier and open microhabitats created by human-induced disturbances. Conversely, they were rare in more natural microsites and microsites with denser vegetation despite their superior persistence ability (polycarpic life cycle). The seed set of tetraploid plants was strongly influenced by their frequency in mixed-ploidy populations. Triploid hybrids originated from bidirectional hybridizations were extremely rare and almost completely sterile, indicating a strong postzygotic barrier between cytotypes. † Conclusions The findings suggest that tetraploids are later immigrants into already established diploid populations and that anthropogenic activities creating open niches favouring propagule introductions were the major factor shaping the non-random distribution and habitat segregation of cytotypes at fine spatial scale. Establishment and spread of tetraploids was further facilitated by their superior persistence through the perennial life cycle. The results highlight the importance of non-adaptive spatio-temporal processes in explaining microhabitat and microspatial segregation of cytotypes.

A taxonomic study of the Pilosella alpicola group growing in the Carpathians revealed the presenc... more A taxonomic study of the Pilosella alpicola group growing in the Carpathians revealed the presence of two morphologically distinguishable taxa: P. ullepitschii (Błocki) Szeląg and P. rhodopea (Griseb.) Szeląg. While P. ullepitschii is endemic to the Carpathians, P. rhodopea is a Balkan subendemic with two isolated localities in the Southern Carpathians (Mt Cozia and Mt Zmeuretu). The core area of distribution of P. ullepitchii is the natural subalpine and alpine meadows of the Western Carpathians (the Vysoké and Západné Tatry Mts in Slovakia and Poland). In addition, only three isolated localities are known from the Nemira Mts (Romanian Eastern Carpathians) and one from the Bucegi Mts (Romanian Southern Carpathians). Interestingly, the Romanian populations occur in man-made habitats (secondary pastures). Karyological and flow cytometric analyses of 305 plants from 13 populations of P. ullepitschii revealed only diploid plants (2n = 2x = 18). One Carpathian population of P. rhodopea from Mt Cozia is also diploid. This is the first report of diploidy in this species. However, the populations from the main part of the distribution of this taxon in the Balkan mountains include other cytotypes. Detailed morphological descriptions and distributions for both taxa are given.

Background and Aims Pilosella officinarum (syn. Hieracium pilosella) is a highly structured speci... more Background and Aims Pilosella officinarum (syn. Hieracium pilosella) is a highly structured species with respect to the ploidy level, with obvious cytogeographic trends. Previous non-collated data indicated a possible differentiation in the frequency of particular ploidy levels in the Czech Republic and Slovakia. Therefore, detailed sampling and ploidy level analyses were assessed to reveal a boundary of common occurrence of tetraploids on one hand and higher ploids on the other. For a better understanding of cytogeographic differentiation of P. officinarum in central Europe, a search was made for a general cytogeographic pattern in Europe based on published data. † Methods DNA-ploidy level and/or chromosome number were identified for 1059 plants using flow cytometry and/or chromosome counting on root meristem preparations. Samples were collected from 336 localities in the Czech Republic, Slovakia and northeastern Hungary. In addition, ploidy levels were determined for plants from 18 localities in Bulgaria, Georgia, Ireland, Italy, Romania and Ukraine. † Key Results Four ploidy levels were found in the studied area with a contrasting pattern of distribution. The most widespread cytotype in the western part of the Czech Republic is tetraploid (4x) reproducing sexually, while the apomictic pentaploids and mostly apomictic hexaploids (5x and 6x, respectively) clearly prevail in Slovakia and the eastern part of the Czech Republic. The boundary between common occurrence of tetraploids and higher ploids is very obvious and represents the geomorphologic boundary between the Bohemian Massif and the Western Carpathians with the adjacent part of Pannonia. Mixed populations consisting of two different ploidy levels were recorded in nearly 11% of localities. A statistically significant difference in a vertical distribution of penta-and hexaploids was observed in the Western Carpathians and the adjacent Pannonian Plain. Hexaploid populations tend to occur at lower elevations (usually below 500 m), while the pentaploid level is more or less evenly distributed up to 1000 m a.s.l. For the first time the heptaploid level (7x) was found on one site in Slovakia. In Europe, the sexual tetraploid level has clearly a sub-Atlantic character of distribution. The plants of higher ploidy level (penta-and hexa-) with mostly apomictic reproduction prevail in the northern part of Scandinavia and the British Isles, the Alps and the Western Carpathians with the adjacent part of Pannonia. A detailed overview of published data shows that extremely rare records on existence of diploid populations in the southwest Alps are with high probability erroneous and most probably refer to the closely related diploid species P. peleteriana. † Conclusions The recent distribution of P. officinarum in Europe is complex and probably reflects the climatic changes during the Pleistocene and consequent postglacial migrations. Probably both penta-and hexaploids arose independently in central Europe (Alps and Carpathian Mountains) and in northern Europe (Scandinavia, Great Britain, Ireland), where the apomictic plants colonized deglaciated areas. We suggest that P. officinarum is in fact an amphidiploid species with a basic tetraploid level, which probably originated from hybridizations of diploid taxa from the section Pilosellina.

Plant Systematics and Evolution, 2012

The genome size of 265 plants and the GC content of 126 plants from 63 populations of the Cyanus ... more The genome size of 265 plants and the GC content of 126 plants from 63 populations of the Cyanus triumfetti and Cyanus montanus groups, collected across the Carpathians, Pannonia, Bohemian Massif, and Western and Dinaric Alps were determined by PI and DAPI flow cytometry. Variation of the nuclear DNA content among homoploid species, and intraspecific and interpopulation variation were confirmed in simultaneous analyses. The 2C-value at the diploid level (the C. triumfetti group) varied from 2.53 for Cyanus dominii subsp. sokolensis to 3.06 pg for C. triumfetti s.s. (1.21-fold range). At the tetraploid level (the C. montanus group), the 2C-value varied from 5.19 for Cyanus mollis to 5.84 pg for C. montanus (1.13-fold range). High intraspecific and interpopulation variation in the amount of nuclear DNA in the C. triumfetti group correlates with the extensive morphological variation found in this group. Significant between-species differences in genome size indicate that this attribute may be used as a supportive taxonomic marker for both of the groups studied. The GC content varied by 2.93 %, from 39.46 % for ''Cyanus axillaris'' to 40.61 % for Cyanus adscendens; this character is of no value for taxonomic purposes. Genome size of the studied populations is significantly higher in southern parts of the distribution area and at higher elevations. Plants with smaller genomes tend to occur in dry areas at low altitudes with high diurnal and annual temperature oscillations. The GC content of the populations studied is significantly correlated with longitude, increasing from east to west; and plants with GC-rich genomes are concentrated in the coldest areas with low minimum temperatures. Keywords Base composition Á Cyanus triumfetti group Á Environmental conditions Á Flow cytometry Á Homoploid species Á Taxonomy Electronic supplementary material The online version of this article (

Plant Systematics and Evolution, 2008

Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian ... more Pilosella alpicola subsp. ullepitschii (Asteraceae) is a strictly allogamous, diploid Carpathian endemic. Its distribution range comprises two areas separated by about 600 km. While in the Western Carpathians (Slovakia and Poland) the taxon occurs in numerous sites, only four localities of man-made origin are known from the Eastern and Southern Carpathians (Romania). We used allozyme markers to test two likely possible scenarios for the origin of this disjunction: long distance dispersal and vicariance. Our data indicate a significant loss of genetic diversity in the isolated Eastern and Southern Carpathian populations in following genetic parameters (averaged per region): percentage of polymorphic loci (38.9% found in the Eastern and Southern Carpathians versus 58.3% in the Western Carpathians), allelic richness (1.4 vs. 1.6), expected heterozygosity (0.134 vs. 0.235), mean number of distinguishable multilocus genotypes (4.3 vs. 10.6) and proportion of distinguishable multilocus genotypes (0.34 vs. 0.68). Higher proportion of homozygous loci found in the Eastern and Southern Carpathian populations might indicate a higher rate of inbreeding due to non-random mating. We assume that these genetically depauperate populations have experienced a very strong genetic bottleneck, probably due to a founder effect. Although our data suggest that the longdistance dispersal model is most likely, more discriminate genetic markers should be used to test this further.

Nordic Journal of Botany, 2007

and Spain are reported. All analysed plants were diploid, with 2n 0 2x010. In addition to P. hier... more and Spain are reported. All analysed plants were diploid, with 2n 0 2x010. In addition to P. hieracioides s.l., we also found diploidy (2n 02x010) in P. nuristanica from Kirgizia and P. japonica from Japan. Our data confirmed previously published results in the literature. We also studied the mode of reproduction in all three taxa. Picris nuristanica and P. japonica s.l. were able to produce some seeds by selfing, and this is the first record of autogamy in this genus. In P. hieracioides s.l. only strict allogamy was found, demonstrating the presence of a self-incompatibility system in this species. Because apomixis, namely apospory, has previously been reported in P. hieracioides, castration experiments were performed in order to confirm or reject this report. However, no evidence of apomictic seed formation was found within this species. In the light of our results, P. hieracioides should be considered a strictly sexual taxon.

Nordic Journal of Botany, 2013

ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically ... more ABSTRACT The Pilosella alpicola group comprises four morphologically distinct and geographically vicariant alpine taxa. We performed a thorough herbarium revision and literature survey to infer their distributional pattern(s). Pilosella alpicola s.s. occurs in the Alps in two disjunct areas: the Swiss Valais Alps and the Italian Dolomites. Historical records come also from the Austrian Alps (Gurktaler Alps and Hohe Tauern) and from one site from the Alpes Maritimes (Col de Larche), but the localities have not been recently confirmed. Pilosella rhodopea, a Balkan subendemic taxon, is quite widespread in Bulgaria (Stara planina Mts, Rila Mts and Pirin Mts), but is more rare in Albania, Greece and Macedonia. Interestingly, this species has also been recorded at two isolated sites in the Romanian southern Carpathians (the Căpăţînii and Cozia Mts). This occurrence underlines the floristic affinities of this part of the Carpathians to the Balkan flora. Only two localities of P. serbica, based on voucher specimens, have been recorded so far; Kopaonik Mts in Serbia and the Prokletije Mts in Montenegro. The records from other ranges are related to P. rhodopea. Pilosella ullepitschii, the detailed distribution of which has already been published, is a Carpathian endemic with its core area of distribution in the western Carpathians (Slovakia and Poland). Three isolated localities are also known in the eastern Carpathians (Nemira Mts) and one locality in the southern Carpathians (Bucegi Mts). The possible causes of disjunctions between and within species ranges are briefly discussed. Based on the distributional data, population sizes and ecology, we evaluate the conservation status of the P. alpicola taxa and propose their inclusion in national Red Lists.