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Chromosome numbers in selected species of Hieracium Sect. Alpina (Asteraceae) from Central and Eastern Europe
Abstract and Figures
Chromosome numbers are reported for 15 taxa ofHieracium sect.Alpina (Griseb.)Gremli from Central and eastern Europe (Austria, the Czech Republic, Poland, Romania, Slovakia, Ukraine). The mode of reproduction
was also studied for some of the taxa. For the first time the chromosome counts of 5 taxa from theHieracium rohacsense group are given:H. rohacsense
Kit. (the West Carpathians),H. ratezaticum (Nyár. etZahn)Mráz (the South Carpathians) and a still unnamed taxon of theH. rohacsense group from Mt. Pop Ivan (the East Carpathians) are tetraploid (2n=36);H. rauzense
Murr (the Eastern Alps) andH. borsanum
Mráz (the East Carpathians) are triploid (2n=27).H. krivanense (Woł. etZahn)Schljakov (the West Carpathians) is tetraploid,H. brevipiliferum
Mráz (the South Carpathians) is triploid; these are the first karyological reports of both taxa belonging to theH. fritzei group. The tetraploid chromosome number was revealed for the first time in an unnamed taxon of theH. nigrescens group from the West Carpathians, and inH. nigrescens subsp.koprovanum
Rech. f. etZahn. InH. alpinum L. s.str. triploid (2n=27) populations from the West Carpathians and diploid populations (2n=18) from the East Carpathians
were confirmed. ForH. halleri
Vill. (theH. alpinum group, the West Carpathians) andH. nigrescens
Willd. (the West Sudeten), the numbers 2n=27 and 2n=36, respectively were found, which is in accordance with previous data. The
triploid level (2n=27) forH. pinetophilum (theH. fritzei group) and the tetraploid level (2n=36) forH. stygium
Uechtr. (theH. chlorocephalum group) both from the West Carpathians were confirmed. One new species, onenomen novum and one new combination at the level of species are published in this paper.
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... There have been relatively few chromosome studies on Hieracium in this region. Chromosome counts have been reported by Skalińska et al. (1959), Skawińska (1963, Májovský (1970a, 1970b, 1974, 1976, 1978), Uhríková & Feráková (1977), Hindáková & Májovský (1977), Mičieta (1978, Murín & Paclová (1979), Hrušovská-Osuská (1988), Chrtek (1996), Szeląg & Jankun (1997), Schuhwerk & Lippert (1999), Májovský et al. (2000) and Mráz (2001b, 2003a, 2003c). Karyotype analysis was given for several previously counted Hieracium taxa by Uhríková (1975) . ...
... Hieracium alpinum group: The counts presented here confirm references from the Western Carpathians (Skalińska in Skalińska et al. 1959, Skawińska 1963, Uhríková & Murín in Májovský 1970b, Murín in Murín & Májovský 1992, Chrtek 1997, Mráz 2001b, Chrtek in Štorchová et al. 2002). The only tetraploid count from the area was published by Szeląg & Jankun (1997), based on plants from Mt Ornak (Tatry Zachodnie Mts, Poland). ...
... The only tetraploid count from the area was published by Szeląg & Jankun (1997), based on plants from Mt Ornak (Tatry Zachodnie Mts, Poland). On the other hand, diploid plants of H. alpinum have been reported from the Eastern Carpathians (Ukraine, Romania) (Chrtek 1997; Mráz 2001b Mráz , 2003c) and from Southern Carpathians (Romania) (Mráz 2003c). In the Carpathians, the diploid and triploid cytotypes are non-overlapping (strictly confined to the eastern, southern and western parts, respectively). ...
Chromosome numbers of 23 species (including subspecies) of Hieracium s. str. from the Western Carpathians are presented. First chromosome numbers are reported for Hieracium kuekenthalianum (=H. tephrosoma, 2n = 36), H. praecurrens (2n = 27) and H. virgicaule (2n = 27); first counts from the Western Carpathians are given for H. atratum (2n = 27), H. bifidum (2n = 27, 36), H, carpathicum (2n = 36), H. inuloides (2n = 27), H. jurassicum (2n = 27), H. macilentum (= H. epimedium, 2n = 27), H. nigritum (2n = 36), H. pilosum (= H. morisianum, 2n=27) and H. silesiacum (2n = 36). New ploidy level (tetraploid, 2n = 36) is reported for H. bupleuroides, hitherto published counts refer only to triploids (2n = 27). Previously published chromosome numbers were confirmed for several other species, i.e. H. alpinum (s.str., 2n = 27), H. bupleuroides (2n = 27), H. crassipedipilum (H. fritzei group, 2n = 27, 36), H. lachenalii (2n = 27), H. murorum (2n = 27), H. prenanthoides (2n = 27), H. racemosum (2n = 27), H. sabaudum (2n = 27), H. slovacum (H. fritzei group, 2n = 36), and H. umbellatum (2n = 18). Triploids and tetraploids predominate, diploids (2n = 18) were found in H. umbellatum. A comprehensive list of previously published chromosome numbers in Hieracium s. str. from the Western Carpathians is provided.
... The counts well support the putative geographic distribution of diploid and triploid cytotypes. While the diploids are most likely confined to the Eastern and Southern Carpathians (Chrtek, 1997;Mráz, 2001;Mráz and Szeląg, 2004), the triploids were repeatedly reported from the remaining parts of the extensive arcto-alpine range of the species (Goldblatt and Johnson, 1979--), incl. the Vranica Mts. , without a precise station). ...
... This is the first chromosome number for the species from Poland, which confirms previously published counts from the Hrubý Jeseník Mts. (Chrtek, 1996) and the Western Carpathians (Chrtek, 1996;Mráz, 2001). ...
Chromosome numbers of 46 Hieracium L. and Pilosella Vaill. taxa from Austria, Bulgaria, Czech Republic, Macedonia, Montenegro, Poland, Romania, Serbia and Slovakia are presented. Chromosomes numbers are given for the first time for Hieracium amphigenum Brig. 2n = 3x = 27, H. bohatschianum Zahn 2n = 4x = 36, H. borbasii R. Uechtr. 2n = 4x = 36, H. cernuum Friv. 2n = 2x = 18, H. hazslinszkyi Pax 2n = 3x = 27, H. mirekii Szelag 2n = 4x = 36, H. polyphyllobasis (Nyar. & Zahn) Szelag 2n = 3x = 27, H. porphyriticum A. Kern. 2n = 4x = 36, H. racemosum Waldst. & Kit. ex Willd. subsp. racemosum 2n = 3x = 27, H. scardicum Borm. & Zahn 2n = 4x = 36, H. sparsum subsp. ipekanum Rech. fil. & Zahn 2n = 4x = 36, H. sparsum subsp. peristeriense Behr & Zahn, H. sparsum subsp. squarrosobracchiatum Behr & al. 2n = 3x = 27, H. tomosense Simk. 2n = 4x = 36, H. tubulare Nyar. 2n = 4x = 36, H. werneri Szelag 2n = 3x = 27 and Pilosella fusca subsp. subpedunculata (Zahn) Szelag, as well as five species of Hieracium sect. Cernua R. Uechtr. not described to date and a hybrid between H. bifidum s. lat. and H. pojoritense Wol.
... Moreover, two of these diploid taxa, namely H. pojoritense (Ștefureac & Tacina, 1979) and the newly described H. vranceae (this paper), are narrow endemics to that region. The third one, a diploid cytotype of H. alpinum, is endemic to both, the Eastern and the Southern Carpathians (Chrtek, 1997;Mráz, 2001;Mráz & Szeląg, 2004;Mráz & al., 2009). Importantly, both H. vranceae and H. pojoritense are strictly associated with relictual calcareous rocky habitats, like the majority of endemic taxa in Europe (Hobohm, 2008;Bruchmann & Hobohm, 2014) or the Carpathians (Kliment & al., 2016;, suggesting their relic character. ...
Hieracium s.str. (Asteraceae) is one of the largest angiosperm genera notorious for its taxonomic complexity caused by widespread interspecific hybridization. This process is tightly coupled with polyploidization and apomixis – asexual reproduction by seeds, which has ensured the persistence of otherwise sterile interspecific hybrids. As a result, apomictic polyploid taxa dominate the taxonomic diversity of the genus whilst sexual diploid species are extremely rare and mostly confined to southern Europe. As diploid taxa are assumed to be the parents of apomictic lineages, the discovery of any new diploid species is important for understanding evolutionary processes and diversity patterns in the genus. Here, we describe a new diploid species, Hieracium vranceae, a narrow endemic to the Munţii Vrancei (Vrancea Mountains, Eastern Carpathians, Romania). This taxon with a distinctive morphology and a strong affinity to relic rocky habitats was first collected in 2013. This means that, even in Europe, some regions, like the Eastern Carpathians, are botanically underexplored and might still preserve an undiscovered diversity of plants. Phylogenetic analyses with multiple molecular markers (low‐copy nuclear genes gsh1 and sqs, nuclear ribosomal ETS, four cpDNA loci) applied to all diploid Hieracium species support specific rank for H. vranceae. In contrast, molecular data suggest conspecifity of two pairs of species, H. lucidum/H. cophanense from Sicily and H. naegelianum/H. renatae from the Balkans, which are moreover only weakly differentiated morphologically. Molecular (cpDNA) and cytogenetic (GISH, FISH) analyses furthermore revealed that H. vranceae has been involved as a maternal parent in the origin of an allotriploid apomictic species, H. telekianum, to which H. vranceae contributed one haplome. The other putative parents of H. telekianum are the diploid narrow endemic H. pojoritense and probably H. sparsum s.l. Our data thus stress the importance of interspecific hybridization for an evolutionary shift from sexuality to apomixis in Hieracium. Our findings furthermore highlight the significance of that part of the Carpathians as one of the most important evolutionary/refugial centres of Hieracium and suggest dynamic species ranges at a regional scale allowing physical contact of taxa whose distributional ranges are nowadays completely allopatric.
... In particular, as diploid maternal plants were considered those which produced diploid embryos by the sexual pathway (ploidy of endosperm:ploidy of embryo 3x:2x), as triploid those which produced triploid embryos by the apomictic pathway (ratio 6x:3x) and, finally, as tetraploid those mother plants which formed tetraploid embryos by the apomictic pathway (ratio 8x:4x). This theoretical assumption was confirmed on selected plants that originated from diploid, triploid and tetraploid taxa (Table 2), when the ploidy of both maternal plant and seeds was determined using conventional flow cytometry or by chromosome counting (cultivated mother plants) as described by Mráz (2001). In addition, the ploidy level, which is uniform within microspecies in Hieracium s.s., was compared with chromosome counts/ploidy level estimations published in numerous literature sources (approx. ...
Apomixis or asexual seed reproduction is a key evolutionary mechanism in certain angiosperms providing them with reproductive assurance and isolation. Nevertheless, the frequency of apomixis is largely unknown, especially in groups with autonomous apomixis such as the diploid–polyploid genus Hieracium.
Using flow cytometric analyses, we determined the ploidy level and reproductive pathways (sexual vs. apomictic) for 7616 seeds originating from 946 plants belonging to >50 taxa sampled at 130 sites across Europe.
Diploid seeds produced by diploids were formed exclusively by the sexual pathway after double fertilization of reduced embryo sacs. An absolute majority of tri- and tetraploid seeds (99.6 %) produced by tri- and tetrapolyploid taxa were formed by autonomous apomixis. Only 20 polyploid seeds (0.4 %) were formed sexually. These seeds, which originated on seven polyploid accessions of four taxa, were formed after fertilization of either unreduced embryo sacs through a so-called triploid bridge or reduced embryo sacs, and frequently resulted in progeny with an increased ploidy. In addition, the formation of seedlings with increased ploidy (4x and 6x) was found in two triploid plants. This is the first firm evidence on functional facultative apomixis in polyploid members of Hieracium sensu stricto (s.s.).
The mode of reproduction in Hieracium s.s. is tightly associated with ploidy. While diploids produce seeds exclusively sexually, polyploids produce seeds by obligate or almost obligate apomixis. Strict apomixis can increase the reproductive assurance and this in turn can increase the colonization ability of apomicts. Nevertheless, our data clearly show that certain polyploid plants are still able to reproduce sexually and contribute to the formation of new cytotypes and genotypes. The finding of functional facultative apomicts is essential for future studies focused on evolution, inheritance and ecological significance of apomixis in this genus.
... Hieracium alpinum comprises both diploid sexual and triploid agamospermous populations, agamospermy has been confirmed by both castration experiments (Chrtek, 1997) and cytoembryological studies (Skawińska, 1963). They are geographically vicariant (diploids in the Eastern and Southern Carpathians, triploids in the remaining parts of the distribution area), population with intermingled diploids and triploids has never been reported (Chrtek, 1997;Mráz, 2001Mráz, , 2003Mráz & Szeląg, 2004). From the Krkonoše Mts, triploid counts were published by Měsíček & Jarolímová (1992), and Chrtek (1994). ...
Five enzyme systems (EST, LAP, PGM, SKD, 6PGDH) were studied in three populations of triploid (2n = 27) agamospermous Hieracium alpinum s. str. (H. alpinum subsp. alpinum) in the Krkonoše Mts (West Sudeten Mts, Czech Republic). Altogether, five different multilocus genotypes were found and both intra-and between-population variation were detected. Within-population variability was found in all the studied populations. Mean population diversity Gsp = 0.52, component of total variance attributed to variation among population Gst = 0.09). Chromosome number 2n = 27 was confirmed in all plants used in enzyme studies. Putative origins of genetic variation are briefly discussed.
Hieracium ciucasense is a new, apomictic species described from the Ciucaş Mountains in the Eastern Carpathians, Romania. It belongs to the H. pietroszense aggregate comprising taxa of presumably hybrid origin between H. alpinum L. and H. bifidum s. lat. The new species is illustrated with photographs of the holotype and living plants; it differs from the other taxa of the H. pietroszense aggregate in the Carpathians in that the leaves are brown-purple spotted. The distribution of the H. pietroszense aggregate in the Carpathians is mapped.
It is for the first time that a research has been done to identify the number of chromosomes and determine their morphology in Hieracium iremelense (Elfstr.) Juxip (Asteraceae), a high-mountain endemic of the central part of the South Urals. It has been found out that the species under investigation is a triploid with 2n=27=3x that often has aneuploid cells with the number of chromosome from 3x+1 to 3x+8. Chromosomes of H. iremelense are medium-sized (2.84±0.04 to 4.82±0.07 µm) and of metacentric and submetacentric types. The variability of morphometric parameters in the chromosomes is characterized by very low and low values of the coefficient of variation. The highest level of variability is observed in the centromeric index and absolute length of chromosomes, the lowest one in their relative length. The revealed karyotypic features of the species under investigation can be used as the material for tackling controversial issues in the systematics of the genus Hieracium L., especially it concerns the species closely related to H. iremelense.
From the aerial parts of Centaurea horrida Bad. (Asteraceae) 25 compounds (two pentacyclic triterpenes, one sterol glucoside, five quinic acid derivatives, one phenolic acid, and 16 flavonoids) were isolated and characterized. The evolutionary meaning of the isolated flavonoids is discussed.
Five isozyme systems (AAT, ADH, LAP, PGM, SKD) were studied in two tetraploid apomictic taxa of the Hieracium rohacsense group (Hieracium sect. Alpina). No intra-and inter-population variation was found in H. rohacsense Kit., endemic to the West Carpathians, which is in accordance with its narrow morphological variation. In contrast, a probably still unnamed tetraploid taxon from Mt. Pop Ivan (Ukrainian East Carpathians) belonging to the H. rohacsense group was represented by three phenotypes detected in one population. The role of diploid sexual taxa occurring in the alpine and subalpine belts of the Ukrainian East Carpathians in maintainig genetic variability is discussed. Apart from morphological characters the two closely related taxa included in this study can be separated also by their patterns of Pgm-1 locus.
A taxonomic concept for the Hieracium nigrescens agg. (H. alpinum ≥ H. murorum) in the Western Carpathians is proposed. Three taxa at the species level are recognized, i.e. Hieracium jarzabczynum, H. mlinicae and H. vapenicanum. One new combination, Hieracium mlinicae (Hruby et Zahn) Chrtek f. et Mráz (H. nigrescens subsp. mlinicae Hruby et Zahn) is published. All taxa should be considered as endemic to the Western Carpathians (both the Polish and Slovakian parts). Detailed descriptions, drawings, lists of localities, distribution maps and determination key are provided along with a comparison with the last comprehensive account of the group (by Zahn 1936). Several lectotypes were chosen for the taxa recognized by Zahn within H. nigrescens s.l.
Taxonomic revision of the Hieracium rohacsense group in the Carpathians revealed that H. rohacsense Kit. is the only representative of this group occuring in the territory of the West Carpathians. The species is endemic to the high mountain ranges of the West Carpathians. A detailed account of distribution is given, with notes on the taxonomy, ecology and phytosociology of the species. Morphological differences from similar taxa co-occurring in the localities of the species are listed.
Mráz, P. & Marhold, K.: Lectotypification of the name Hieracium rohacsense Kit. (Compositae). — Willdenowia 29: 41–44. 1999. — ISSN 0511-9618. A lectotype is designated for Hieracium rohacsense, the name of a species restricted in its distribution to the subalpine belt of the W Carpathians with two doubtful localities in the E Carpathians. The lectotype is a specimen collected in the Západné Tatry (Roháce) Mts in Slovakia and kept in the Kitaibel herbarium in the Hungarian Museum of Natural History (BP) in Budapest.
Hieracium decipientiforme (Wołoszczak et Zahn) Šljakov, most probably an endemic taxon of the Ukrainian Carpathians, was found to be agamospermous tetraploid (2n=36). It occurs in mountain meadows, open places in krummholz stands and in subalpine grasslands in altitudes 1580-1900 m a.s.1. in the Gorgany Mts, the Čornohora Mts and the Marmaros Mts. Important distinguishing characters and relations to other related species groups are briefly discussed.
Chromosome numbers are reported for 18 collections representing ten taxa of Hieracium L. Chromosome numbers are reported for the first time for H. chlorocephalum Uechtr. (2n = 4x = 36), H. stygium Uechtr. (2n = 36), H. silesiacum Krause (2n = 36), H. corconticum Knaf fil. ex Čelak. (2n = 27), H. conicum Arvet-Touvet (2n = 18), and H. chrysostyloides (Zahn) Chrtek jun. (2n = 5x = 45). The pentaploid chromosome number appears to be only the second count for any taxon of Hieracium s. str. above the level of tetraploid. In H. prenanthoides Vill. 2n = 27 was ascertained in populations from the Krkonoše Mts. (West Sudeten Mts.) and Vysoké Tatry Mts. (Western Carpathians). The mountain population of H. schmidtii Tausch from the Krkonoše Mts. (West Sudeten Mts.) was found to be triploid (2n = 27). In H. transsilvanicum Heuff. the number 2n = 18 was confirmed, and in H. nigritum Uechtr. 2n = 36 was ascertained.
“Apomixis” in angiosperms means asexual (agamic) reproduction by seeds, i.e, “agamospermy” (“seed apomixis”). The normal counterpart of this widespread anomaly of reproduction is “amphimixis”, i.e., sexual reproduction. Orginally, Winkler (1908) defined apomixis in a wider sense, including vivipary, pseudovivipary, and other forms of vegetative propagation such as bulbils, runners, rhizomes, etc. This broader definition is now rather rarely used.
The mapping scheme for the Flora of Central Europe forms the basis for the preparation of a distributional atlas. This atlas should contain the maps of all Central European vascular plants (Pteridophyta and Spermatophyta) based on grid units of 10′ long. × 6′ lat. The maps will cover Austria, western and central Czechoslovakia, Germany (G.D.R. and G.F.R.), Switzerland, northern Italy and northwestern Yugoslavia (north of 45° N and west of 19° 10′ E), together with some neighbouring territories.
Work on the mapping scheme was started a few years ago in most Central European countries with the help of numerous voluntary collaborators, many local working groups, and several regional centers. The current activity is characterized by international cooperation and the accumulation of data prepared for modern machine handling. A “Check list of the vascular plants of Central Europe” (Ehrendorfer & al., cf. fig. 1) and a grid system based on geographical longitudes and latidudes supply the numerically coded taxonomical and geographical reference units. The system of data handling is demonstrated in fig. 2: Records from field work as well as herbarium and literature sources are first incorporated into printed field lists (fig. 3), punch cards with text (fig. 4) or data forms (fig. 5). Records are then transferred to punched cards or magnetic tape, partly through mark‐sensing coding‐forms and mark‐sensing reader. Finally, lists of records as well as distributional maps will be available by automatic printing from tabulators and/or data plotters.
10 sample maps demonstrate the design and information value of the distributional maps for the atlas; at the same time they illustrate the considerable deficiencies in our present knowledge of actual distribution in higher plants. Everybody interested in Central European floristics and geobotany is therefore invited to participate in regional or national organizations of the mapping scheme in order to realize the project of a distributional atlas of the Central European flora.
Chromosome numbers are given of 15 species of the genus Hieracium L. s. str., representing seven species groups (in the sense of Flora Europaea, roughly corresponding to Zahn's "species principales") from the Krkonoše Mts., N. Bohemia and SW Poland. For the first time, chromosome numbers are reported for H. melanocephalum Tausch (2n=27), H. tubulosum Tausch (2n=36), H. schustleri Zlatník (2n=36), H. fritzei F. Schultz (2n=27), H. rohlenae Zlatník (2n=27), H. nigrescens Willd. (2n=36), H. decipiens Tausch (2n=36), H. atrellum Juxip in Schischkin et Bobrov (2n=27), H. subnigrescens (Fries ex Norrlin) Dahlst. (2n=36), H. sudeticum Sternberg (2n=36), H. pedunculare Tausch (2n=36), H. glandulosodentatum Uechtr. (2n=36), H. wimmeri Uechtr. (2n=27). In Hieracium alpinum L. s. str. the number 2n=27 has been confirmed. The results show a high proportion of tetraploid taxa; no diploids have been found.
