Chromosome numbers and DNA ploidy levels of selected species ofHieracium s.str. (Asteraceae)

Chromosome numbers and /or ploidy levels are reported for 44 species and subspecies ofHieracium s.str. from the following European countries: Andorra, Austria, Bulgaria, Czech Republic, France, Italy, Montenegro, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland and Ukraine. The chromosome numbers/DNA ploidy levels ofH. bocconei (2n ～ 4x),H. bupleuroides subsp.leviceps (2n = 27),H. caesioides subsp.caesioides (2n = 27),H. basifolium (H. caesium agg., 2n = 36),H. plumbeum (H. caesium agg., 2n = 36),H. glaucum subsp.nipholepium (2n= 27),H.gouanii (2n = 18),H. gymnocerinthe (2n = 27),H. ramondii (2n = 27),H. recoderi (2n = 18),H. stelligerum (2n = 18), andH. tomentosum (2n = 18, 2n ～ 2x, 2n ～ 3x) were determined for the first time. New ploidy levels are reported forH. cerinthoides s.str. (2n = 27),H. humile (2n = 36), andH. tommasinianum (2n = 27).     

Chromosome number variability in central european members of the<Emphasis Type="Italic">Festuca ovina</Emphasis> and<Emphasis Type="Italic">F. pallens</Emphasis> groups (sect.<Emphasis Type="Italic">Festuca</Emphasis>)

Chromosome numbers for 98 plants ofF. pallens, 19 ofF. psammophila, F. belensis andF. vaginata, and 44 ofF. ovina (originating from Austria, the Czech Republic, Germany, Slovakia and Latvia) are given. In addition to theF. ovina andF. pallens groups, chromosome counts for the following taxa are also reported:F. alpestris (2n=14) reported for the first time in this work,F. amethystina subsp.amethystina (2n=28),F. brevipila (2n=42),F. cinerea (2n=28),F. rupicola subsp.rupicola (2n=42) andF. versicolor subsp.versicolor (2n=14).InF. pallens, two ploidy levels (2n=2x=14+0-1B, 2n=4x=28+0-1B) as well as two natural triploid plants (2n=21+0-1B), were found. In addition to the fourF. pallens types that have been distinguished in Austria, one new tetraploid type (F. pallens “scabrifolia”) from the Czech Republic and Germany is reported and its taxonomy is discussed. The distributions of the Oberösterreich-Niederösterreich and Pannonisches-HügellandF. pallens types outside of Austria are documented.Only the diploid chromosome number (2n=14) was found inF. psammophila andF. vaginata. Chromosome numbers forF. psammophila subsp.muellerstollii andF. belensis (both 2n=14) were determined here for the first time. Two ploidy levels, 2n=14+0-5B corresponding toF. ovina subsp.ovina and 2n=28 corresponding toF. ovina subsp.guestphalica andF. cf.duernsteinensis were confirmed inF. ovina. Differences in chromosome structure (simple and multiple secondary constrictions) betweenF. pallens as opposed toF. psammophila andF. vaginata are discussed. A complete survey of published chromosome counts for Central European species from theF. ovina andF. pallens groups is included.     

Observations of meiotic chromosomes in Gaura (Onagraceae)

Observations of meiotic chromosomes are reported for all 21 species and 3 additional sub species ofGaura (Onagraceae), based upon a study of 647 individuals from 509 naturally occurring populations throughout the range of the genus. The basic chromosome number for the genus isx = 7, and 18 species are diploid withn = 7. Among these, the self-incompatible ones are often highly chromosomally heterozygous, with no homozygous individuals having been found in nature in the perenrennialsGaura lindheimeri andG. villosa, and two-thirds or more of the individuals apparently heterozygous in the following well-sampled species:G. calcicola, G. longiflora, andG. suffulta subsp.suffulta. In contrast, the autogamous species are entirely chromosomally homozygous or nearly so. Two species ofGaura are reported as chromosomal structural heterozygotes, with about 50% pollen abortion:G. biennis andG. triangulata; the translocation systems originated independently of one another. Two of the three polyploid species,G. sinuata andG. drummondii (G. odorata of many authors), are consistently tetraploid (n = 14) and, despite their cytological autotetraploidy, are thought to have originated following interspecific hybridization. They are the only rhizomatous species in the genus and may have had one ancestor in common. The remaining polyploid,G. coccinea, includes populations withn = 7, 14, 21, and 28, as well as evident interploid hybrids and, frequently, supernumerary chromosomes. The relationship among these populations is close and is maintained by frequent hybridization and exchange of genetic material. No other species seems to have participated in their origin, and the association of their chromosomes is consistently that characteristic of autopolyploidy in plants with tetraploid and higher chromosome numbers.     

Karyological studies in IberianSonchus (Asteraceae: Lactuceae): S. oleraceus, S. microcephalus andS. asper and a general discussion

Chromosome numbers and karyotypes ofSonchus oleraceus (n=16, 2n=32),S. asper (n=9, 2n=18 for subsp.asper and subsp.glaucescens, andS. microcephalus (n=15, 2n=30) are studied; for the first time their idiograms are given. Karyotypes mainly comprise small chromosomes with a degree of asymmetry 2B inS. oleraceus andS. microcephalus and 2A or 2A-2B inS. asper. No karyological differences can be pointed out betweenS. asper subspecies. Data presented support the amphiploid character ofS. oleraceus, and the origin ofS. microcephalus through a dysploid process involving the former taxon. In Iberian representatives of the genus, diagrams of karyotype asymmetry indices show a cluster grouping for species, with the detached exceptions ofS. maritimus andS. crassifolius, which spontaneously hybridize in central Spain. A review of available karyological data shows that in the evolution of the genusSonchus s.l. and relatives, the basic chromosome number x=9 has generally been maintained. Dysploidy is restricted to the seriesS. bourgeaui (n=8) —S. tenerrimus (n=7) andS. oleraceus (n=16) —S. microcephalus (n=15), evolutionarily related and included in the present subgenusSonchus. Polyploidy has been detected in a total of nine taxa ofSonchus and in the generaEmbergeria, Kirkianella, andDendroseris, being more common in peripheral regions of the distribution area of the group. Five diversification centers are proposed for the whole group, of which the Western Mediterranean area, including the Iberian Peninsula, is related to diversification of the present subgenusSonchus.     

New or rare data on chromosome numbers in several taxa of the genus<Emphasis Type="Italic">Artemisia (Asteraceae)</Emphasis> in Poland

Chromosome counts in 16 populations of fiveArtemisia species from Poland are presented in this paper. Those ofA. annua (2n=18) andA. dracunculus (2n=90) are reported for the first time in Polish populations. The decaploid level (2n=90) is described for the first time in non-cultivated populations ofA. dracunculus, and several cases of aneusomaty (intraindividual aneuploid variations in chromosome number: 2n=87, 88 and 89) have been detected in this species. In addition to the already known diploid number (2n=18), the tetraploid level (2n=36) has been detected inA. absinthium. The same two numbers have been recorded inA. abrotanum, which represents the first tetraploid count in populations of this taxon occurring outside botanical gardens. Finally, the chromosome number ofArtemisia campestris subsp.sericea (tetraploid, 2n=36) is reported for the first time. The relevance of polyploidy for the evolution of the genus and other cytotaxonomic or cytobiogeographical aspects are briefly discussed.     

Chromosome numbers within the genusBolboschoenus in Central Europe

Two chromosome numbers n=54, n=55 were found inBolboschoenus plants studied from Central Europe (Czech Republic, Slovakia, Hungary, Poland) and coastal regions of Europe (the Netherlands, Sweden). The number n=55 is typical forB. maritimus subsp.maritimus with narrow fruits and mostly also forB. maritimus subsp.compactus; the number n=54 characterizesB. planiculmis auct. The morphological type ofB. maritimus subsp.maritimus with wide fruits represents a stable taxon occurring in freshwater habitats throughout Europe. Its variation in chromosome numbers (both n=54, n=55) indicates a possible hybrid origin, probably resulting from hybridization betweenB. maritimus subsp.maritimus with narrow fruits andB. planiculmis auct. Spontaneous hybridization betweenBolboschoenus taxa in the regions with mixed populations may explain the origin of the intermediate morphological and anatomical characters of plants from some localities and the deviations in chromosome numbers.     

Ploidy variation in the group ofMyosotis palustris andM. laxa in the Czech Republic and Slovakia

In 74 natural populations of theMyosotis palustris andM. laxa groups from former Czechoslovakia three euploid and one aneuploid cytotypes were found that belong to the following species:M. nemorosa Besser, 2n=22,M. palustris (L.)Nath. em.Reichenb. inSturm, 2n=66,M. laxiflora Reichenb., 2n=66,M. radicans Opiz, 2n=66(64),M. brevisetacea (Schuster) Holub, 2n=66(64), andM. caespitosa C.F. Schultz, 2n=88. With the exception ofM. nemorosa, the chromosome numbers are published for the first time for the Czech Republic. The chromsome numbers, 2n=66, 2n=64 are found inM. brevisetacea, are new for the literature. The correlation between pollen size and ploidy level has been studied as well, and statistically significant differences were found between cytotypes belonging to different ploidy levels.     

The annual species ofAdonis (Ranunculaceae)—a polyploid complex

The five annual species ofAdonis L., sect.Adonis, growing in Israel, form a series of diploid, tetraploid and hexaploid species. Their somatic chromosome numbers are 2n = 16 inA. annua L.,A. dentata Del. andA. palaestina Boiss., 2n = 32 inA. microcarpa DC., 2n = 48 inA. aestivalis L.; counts forA. dentata, A. palaestina andA. microcarpa are new records. There are indications that alloploidization may have been involved in the process of speciation in sect.Adonis. A taxonomic survey of the 8 species of the section reveals that a higher ploidy level is usually combined with a larger distribution area.     

Karyotypes of three species of the family Cottidae (Scorpaeniformes)

Karyotypes and cellular DNA contents of three species of the family Cottidae viz.Icelus cataphractus, Gymnocanthus intermedius andAlcichthys alcicornis were analyzed. Structural modifications within the family were supposedly by Robertsonian translocations. The diploid chromosome numbers were determined to be 48 inAlcichthys alcicornis, 44 inGymnocanthus intermedius and 40 inIcelus cataphractus. The DNA contents ranged from 1.46 to 1.50pg/cell in the three species. The karyotype ofIcelus cataphractus is unique in having the smallest chromosome number (2n = 40) and 14 large-sized chromosomes. From the chromosome number and the existence of some large chromosome pairs, Robertsonian translocations seem to have occurred frequently inIcelus cataphractus andGymnocanthus intermedius.     

Variation of Giemsa C-band and fluorochrome banded karyotypes,and relationships inAllium subgen.Molium

The somatic karyotypes of 10 taxa belonging toAllium subgen.Molium (Liliaceae) from the Mediterranean area have been investigated using Giemsa C-band and fluorochrome (Hoechst, Quinacrine) banding techniques. A wide range of banding patterns has been revealed. InAllium moly (2n = 14),A. oreophilum (2n = 16) andA. paradoxum (2n = 16) C-banding is restricted to a region on each side of the nucleolar organisers and the satellites show reduced fluorescence with fluorochromes. The satellites are also C-banded and with reduced fluorescence inA. triquetrum (2n = 18), but two other chromosome pairs also have telomeric bands which are not distinguished by fluorochrome treatment. InA. erdelii (2n = 16) 4 pairs of metacentric chromosomes have telomeric C-bands while 2 pairs of telocentric chromosomes have centromeric C-banding. InA. subhirsutum (2n = 14),A. neapolitanum (2n = 28),A. trifoliatum subsp.hirsutum (2n = 14) andA. trifoliatum subsp.trifoliatum (2n = 21) chromosomes with long centromeres, consisting of a centromere and nucleolar organiser are positively C-banded on each side of the constriction. InA. subhirsutum banding is confined to the pair of chromosomes with this feature, whereas inA. neapolitanum one additional chromosome pair has telomeric bands and inA. trifoliatum there are varying numbers of chromosomes with centromeric and telomeric bands, depending on the subspecies.A. zebdanense (2n = 18) shows no C-bands. The banding patterns in this subgenus are compared with those recorded for otherAllium species and with the sectional divisions in the genus. Evidence from the banding patterns for allopolyploidy inA. trifoliatum subsp.trifoliatum andA. neapolitanum is discussed.     

Karyotype evolution by centromeric fission inZamia (Cycadales)

The chromosome numbers of several species ofZamia from Mexico are reported.Z. paucijuga, distributed from central Oaxaca to Nayarit, has been found to have 2n = 23, 25, 26, 27 and 28. 2n = 28 is the highest chromosome number yet found in the cycads. Karyotypes of this species differ principally in the number of telocentric and metacentric chromosomes present in each; 2n = 23, 25, 26, 27 and 28 were found to have 5, 3, 2, 1 and 0 metacentric and 8, 12, 14, 16 and 18 telocentric chromosomes, respectively.Z. fischeri has been found to be 2n = 16,Z. furfuracea andZ. loddigesii 2n = 18.Zamia paucijuga on the basis of morphological and ecological characteristics, is considered to be an advanced member of this genus. Chromosome and karyotype evolution inZ. paucijuga may have occurred by centromeric fission of metacentric chromosomes; the karyotypes ofZ. paucijuga are strongly asymmetrical, suggesting that they evolved recently.     

Karyological studies onGentiana sect.Chondrophyllae (Gentianaceae) from China

Nineteen populations of fifteen species ofGentiana sect.Chondrophyllae from China were observed cytologically.Gentiana alsinoides, G. anisostemon, G. asterocalyx, G. exigua, G. heterostemon, G. intricata, G. praticola, G. pseudoaquatica, G. spathulifolia, andG. subintricata all had the same chromosome number of 2n = 20 (or n = 10), whereasG. piasezkii had 2n = 36,G. squarrosa 2n = 38,G. prattii 2n = 18,G. aristata 2n = 14 (n = 7), andG. heleonastes 2n = 12. All these chromosome numbers are documented here for the first time, except forG. squarrosa, where it is a new number report. The basic numbers of x = 6, x = 7 and x = 19 are new for the section. Karyotype analyses of some species revealed that, except for a few cases, the species examined mainly had metacentric chromosomes. 2n = 20 = 2m(SAT) + 18m was found to be the main type of karyotype for the species with 2n = 20. Chromosomal evolution and its mechanism in this section are also discussed.     

Comparative karyomorphology and interrelationships ofSelaginella in Japan

Karyomorphological comparisons were made of 16 native and cultivated species ofSelaginella in Japan. The somatic chromosome numbers are 2n=16 inS. boninensis; 2n=18 inS. doederleinii, S. helvetica, S. limbata, S. lutchuensis, S. nipponica, S. selaginoides, S. tama-montana, andS. uncinata; 2n=20 inS. biformis, S. involvens, S. moellendorffii, S. remotifolia, andS. tamariscina; 2n=30 inS. rossii; and 2n=32 inS. heterostachys. The interphase nuclei of all species examined are uniformly assigned to the simple chromocenter type. The metaphase karyotype of 2n=16 (x=8) is 8 m (=median centromeric chromosomes)+8(st+t)(=subterminal and terminal). The group of the species having 2n=18 (x=9) is heterogeneous karyomorphologically: The karyotype ofS. nipponica is 2n=18=6 m+12(st+t),S. tama-montana 10 m+2 sm(=submedian)+6(st+t), andS. uncinata 6 m+7 sm+5(st+t). Although the remaining five species have the common karyotype 8 m+4 sm+6(st+t), the values of mean chromosome length are variable. Another group of the specles having 2n=20 (x=10) is homogeneous, since all species have the same karyotypes 8 m+4 sm+8(st+t) and have similar chromosome size. The karyotype of 2n=30 is 12 m+6 sm+12(st+t) and is suggested to be a triploid of x=10, and 2n=32=16m+16(st+t), a tetraploid of x=8. Thus, three kinds of basic chromosome numbers, x=8, 9, 10 are present in JapaneseSelaginella examined, and their karyomorphological relationships are discussed.     

Genome size variation and C-band polymorphism inAlstroemeria aurea, A. ligtu andA. magnifica (Alstroemeriaceae)

Among a total of 43 accessions ofAlstroemeria aurea, A. ligtu andA. magnifica nuclear DNA amounts (2C-values) showed significant intraspecific variation, 1.09, 1.21 and 1.15 fold, respectively, when determined through flow cytometric measurements of fluorescence of propidium iodide (PI) stained nuclei. After staining with another fluorochrome, 4,6-diamidino-2-phenylindole (DAPI), an intraspecific variation of 1.10, 1.11 and 1.12 fold, respectively, was found. C-band polymorphisms were present among and within the accessions of all three species. In some cases only very small differences in C-banding pattern were observed. In other cases, however, differences were more prominent. Besides C-band polymorphism, there were also instances of chromosome length polymorphism, which concerned the total chromosome complement or single chromosomes. The variation in nuclear DNA amount inA. aurea andA. ligtu was more or less continuous, except for one accession ofA. ligtu subsp.simsii. Artificial selection and possibly introgression of chromosomes from other species may have moulded the karyotypes of some of the accessions ofA. aurea, a species that has been under cultivation for more than 160 years. The variation as observed inA. magnifica subsp.magnifica was discontinuous and could be due to a broad species concept.     

The allopolyploid origin ofSedum rupestre subsp.rupestre (Crassulaceae)

InSedum rupestre L. a polyploid series (x = 16) occurs in which aneuploid chromosome numbers and odd levels of ploidy prevail. The most common and widely distributed cytotype,S. rupestre subsp.rupestre, is 2n = 112. Plants resemblingS. rupestre subsp.rupestre can be obtained by hybridizing the tetraploid cytotypes ofS. forsterianum Sm. (2n = 48) andS. rupestre subsp.erectum 't Hart (2n = 64). Comparison of these artificial hybrids with their parents and a large number of plants ofS. rupestre subsp.rupestre (2n = 112) from nature showed thatS. rupestre subsp.rupestre and the artificial hybrids are morphologically indistinguishable, and intermediate betweenS. forsterianum andS. rupestre subsp.erectum. MorphologicallyS. rupestre subsp.rupestre is closer to subsp.erectum than toS. forsterianum. Chloroplast DNA restriction patterns ofS. rupestre subsp.rupestre, however, resembleS. forsterianum more closely. The combined results of the hybridization experiments, the analysis of the cpDNA restriction patterns, and the morphological variation indicate the allopolyploid origin ofS. rupestre subsp.rupestre. Natural hybrids inSedum (Crassulaceae) 4.     

Polyploidy and aneuploidy inOphrys,Orchis, andAnacamptis (Orchidaceae)

Studies on chromosome numbers and karyotypes in Orchid taxa from Apulia (Italy) revealed triploid complements inOphrys tenthredinifera andOrchis italica. InO. tenthredinifera there is no significant difference between the diploid and the triploid karyotypes. The tetraploid cytotype ofAnacamptis pyramidalis forms 36 bivalents during metaphase I in embryo sac mother cells. Aneuploidy was noticed inOphrys bertolonii ×O. tarentina with chromosome numbers n = 19 and 2n = 38. There were diploid (2n = 2x = 36), tetraploid (2n = 4x = 72), hexaploid (2n = 6x = 108) and octoploid (2n = 8x = 144) cells in the ovary wall of the diploid hybridOphrys apulica ×O. bombyliflora. Evolutionary trends inOphrys andOrchis chromosomes are discussed.     

Chromosomal evolution withinPiperaceae

Chromosome numbers for 26 different species of the generaPiper, Peperomia andPothomorphe (Piperaceae) are reported. The basic chromosome numbers are 2n = 26, x = 13 (Piper, Pothomorphe) and 2n = 22, x = 11 (Peperomia), polyploid series are characteristic forPiper andPeperomia. Piper has the smallest chromosomes and prochromosomal interphase nuclei,Peperomia the largest ones and mostly reticulate to euchromatic nuclei.Pothomorphe is intermediate in both characters. The karyomorphological differences betweenPothomorphe andPiper underline their generic separation. Interspecific size variation of chromosomes occurs inPiper andPeperomia. Infraspecific polyploidy was observed inPiper betle. C-banding reveals different patterns of heterochromatin (hc) distribution between the genera investigated. The genome evolution is discussed.     

The karyology of some neotropicalStyracaceae

Chromosome numbers and karyomorphological characters have been investigated inPamphilia andStyrax (Styracaceae). Counted for the first time in the genus, two species ofPamphilia were found to have 2n = 16. The twoStyrax spp. investigated share withPamphilia the same chromosome number, a peculiar condensation behaviour of the chromosomes (Fig. 1a–c) and the same type of semi-reticulate interphase nucleus, results which indicate a close relationship of the two genera. The base number inStyracaceae is probably x = 8 (2n = 2x = 16) with stabilized triploids inHalesia andPterostyrax (2n = 3x = 24). A preliminary comparison withSapotaceae andEbenaceae does not allow a general karyological characterisation of the orderEbenales.     

Taxonomic studies ofAsarum sensu lato

The karyotype and the C-banding pattern in two species ofHexastylis andAsarum epigynum were analysed in detail, and the results obtained were compared with those of the other species ofAsarum, Asiasarum andHeterotropa previously reported. The present results were partially different from the previous reports related to the karyotypes of these species. The karyotype observed in two species ofHexastylis (2n=26) was represented by ten pairs of metacentric chromosomes and three pairs of small subtelocentric chromosomes, which is very similar to that ofAsiasarum in eastern Asia. The C-banding patterns ofHexastylis andAsiasarum, however, were clearly different from each other. A striking difference was found in one of the three pairs of small subtelocentric chromosomes. A Formosan speciesAsarum epigynum had the somatic chromosome number 2n=12 and a highly asymmetrical karyotype composed of mainly subtelocentric chromosomes. These karyological features were remarkably different from those of the other groups inAsarum s.l.     

A biosystematic study on polyploid populations of the genusSpiraea (Rosaceae) in Korea

A total of 61 chromosome counts from 11 taxa of Korean Spiraea was made. Our counts are the first report for eight taxa;S. blumei (2n=18, 36),S. chartacea (2n=36),S. chinensis (2n=36),S. fritschiana (2n=27, 36),S. microgyna (2n=18),S. prunifolia var.simpliciflora (2n=18),S. pseudocrenata (2n=36), andS. trichocarpa (2n=18). A new chromosome number of 2n=36 (tetraploid) is reported forS. pubescence. Populations of three species includingS. blumei, S. pubescence and .V.fritschiana, show different ploidy levels; diploid and tetraploid populations are found in the former two species and triploid and tetraploid ones, in latter species. Multiplication of chromosome numbers contributes to increase in size of pollen and stomata in the three species. Populations with different ploidy levels inS. blumei occupy different regions; diploid populations in inland Korea and tetraploid ones in Ullung Island. Island tetraploid population of 5.blumei might be originated from intra-island polyploidization through the introduction of diploid from inland Korea, considering the worldwide distribution of this species. Pollen fertilities of island populations of ,S′.blumei are relatively low, and sometimes no pollen grain is produced in anther sacs; it suggests that tetraploid population of the island is gynodioecious which may serve reduction of inbreeding depression.