Cytogenetic studies on natural intergeneric hybridization inAster alliances

Intergeneric hybrids ofAster ageratoides subsp.ovatus (2n=36)×Kalimeris pinnatifida (2n=18) were produced artificially. The chromosomes of the hybrid were found to be 2n=27, and to consist of 9 large chromosomes and 18 small chromosomes. In meiosis of the PMCs of the hybrid, a chromosome configuration of 9_II+9_I was regularly observed. While all the univalents were large, and all the bivalents were comparatively small. The large and small chromosomes ofA. ageratoides subsp.ovatus were found to be rather distant in homology, and the small chromosomes of the subspecies and the chromosomes ofK. pinnatifida were found to have a high degree of homology. The tetraploidovatus was concluded to be an amphidiploid, composed of the large chromosomes ofAster and the small chromosomes ofKalimeris.     

CHROMOSOME NUMBER DETERMINATIONS IN ASTER L. (COMPOSITAE) WITH COMMENTS ON CYTOGEOGRAPHY,PHYLOGENY AND CHROMOSOME MORPHOLOGY

Chromosome number determinations from 360 individuals of 73 species, varieties and hybrids of Aster (not including Virgulus Raf.) are reported for the first time. Most reports confirm one or more earlier ones for the taxa. Several first counts are included: A. curtisii T. & G., 2n = 32; A. drummondii Lindl. var. drummondii, 2n = 16; A. eulae Shinners, 2n = 48; A. infirmus Michx., 2n = 18; A. lateriflorus (L.) Britt., 2n = 64; A. meritus A. Nels., 2n = 36; A. parviceps (Burg.) Macken. & Bush, 2n = 32; A. pubentior Cronq., 2n = 18; and A. sagittifolius Wedem., 2n = 48. Counts of 2n = 18 for A. avitus Alex. and A. hemisphaericus Alex. and 2n = 36 for A. paludosus Ait. provide additional evidence that the base number of A. sect. Heleastrum is x = 9, not x = 5. Counts of 2n = 32 and 2n = 48 for A. curtisii support its placement with other x = 8 species in sect. Foliacei, but not with x = 9 species in sect. Spectabiles. The geographic distributions of cytotypes support the taxonomic delimitations A. puniceus L. and A. praealtus Poir. Diploids were found throughout the range of A. puniceus, but tetraploids were rare and found only in the southwestern portion of the range; ploidy level did not correlate with morphological variation. In A. praealtus the distribution of tetraploids and octoploids correlated with the allopatric varities praealtus and angustior Wieg. Satellite chromosome morphology was determined in species not studied before. Aster engelmannii, A. glaucodes, A. infirmus, A. modestus, A. oregonensis, A. pauciflorus and A. pubentior had subequal satellites and short arms, the “primitive” type in the genus. Aster meritus, A. radula and A. radulina had the “derived,” subg. Aster type, a very large satellite attached to a very short proximal portion of the short arm.     

Cytogenetic studies on natural intergeneric hybridization inAster alliances

In addition to various types of natural hybrids betweenAster ageratoides subsp.ovatus andKalimeris incisa reported earlier, a new backcross type has been discovered. This new type, characterized by the chromosomes of 2n=27L+54S, was most probably produced through fertilization of a normally-reduced gamete of the F₁ plant (2n=72=18L+54S) and an unreduced gamete of subsp.ovatus (2n=36=18L+18S).     

KARYOTYPIC VARIATION OF EASTERN NORTH AMERICAN CLAYTONIA CHEMICAL RACES

Considerable karyotypic differentiation has occurred within the group of taxa comprising the eastern North American members of the genus Claytonia. Patterns of karyotypic differentiation are congruent with evolutionary groupings based on flavonoid chemistry, particularly at the diploid level. The 2n = 16 diploid chemotype found in both C. caroliniana and C. virginica possesses a karyotype composed entirely of metacentric chromosomes, while acrocentric chromosomes predominate in the karyotypes of the 2n = 12 and 2n = 14 diploid chemotypes of C. virginica. The 2n = 16 diploid also has a karyotype significantly larger than those of the other diploid cytotypes. Polyploid karyotypes of both species show varying degrees of divergence from their presumed diploid progenitors.     

AMPHIPLOID ORIGIN OF TWO ENDEMIC RACES OF ASTER (ASTERACEAE) IN SOUTHERN CALIFORNIA

Two cytological races of Aster which morphologically resemble A. ascendens but differ from it in chromosome number are found in the Transverse and Peninsular ranges of southern California. One of these races is relatively common, can usually be separated morphologically from A. ascendens, and has a somatic chromosome number of 36; it corresponds to the endemic species A. bernardinus. The other race is rare and localized, is morphologically almost indistinguishable from A. ascendens, and has a somatic chromosome number of 42. The chromosome numbers and karyotypes of the two races suggest that they originated through backcrosses of A. ascendens (x = 13) to its respective parent taxa, A. falcatus (x = 5) and A. occidentalis (x = 8).     

Comparison of the Giemsa C-banded karyotypes of the three subspecies ofPsathyrostachys fragilis,subspp.villosus (2x),secaliformis (2x, 4x), andfragilis (2x) (Poaceae), with notes on chromosome pairing

The karyotypes of diploidP. fragilis subsp.villosus (2n = 2x = 14) and tetraploid subsp.secaliformis (2n = 4x = 28) were studied by Giemsa C- and N-banding, and AgNO₃ staining and compared with the karyotype of subsp.fragilis (2x). The complements of subsp.villosus and subsp.fragilis were similar, with 8 metacentric and 6 SAT-chromosomes, one metacentric and two submetacentric pairs, with small to minute, polymorphic, heterochromatic satellites. The complement of subsp.secaliformis on the whole agreed with a doubling of the complement of diploidP. fragilis, suggesting autopolyploidy. Only the presence of 12 nucleoli in interphases identified 6 SAT-chromosome pairs. In subsp.villosus one or two extra micronucleoli indicated a chromosome pair with very low nucleolusforming activity, bringing the number of SAT-chromosome pairs to 4. This number may be a characteristc ofPsathyrostachys. Besides very small, inconsistently observed bands, the C-banding pattern consisted of 0–3 small bands per chromosome at intercalary and terminal locations, and at NORs. The level of banding pattern polymorphism was low, but enough to indicate that the taxa are outbreeders. Similarities in chromosome morphology and C-banding patterns identified homology of all chromosomes of subsp.villosus, but for 12 pairs only in subsp.secaliformis. Between plants, reliable identification of homology and homoeology (subsp.secaliformis) was possible only for the SAT-chromosomes and the shortest metacentrics. Chromocentres were very small and the amount of constitutive heterochromatin was low. N-banding stained chromosomes uniformly. The basic karyotypes of theP. fragilis taxa were similar to those ofP. juncea, P. lanuginosa, andP. stoloniformis supporting a close relationship and the presence of a common genome, N. NORs had different nucleolus-forming activities. Meiotic analysis demonstrated a high level of bivalent pairing in the three taxa. A chromosomal rearrangement was suggested in subsp.villosus. The low multivalent frequency in subsp.secaliformis indicates the presence of a pairing regulation mechanism. The majority of chiasmata were interstitial. Pollen grain size discriminated between diploid and tetraploid taxa. The existence of a diploid cytotype of subsp.secaliformis is supported by pollen measurements of herbarium material.     

Extensive chromosome number variation in Aster ageratoides var. pendulus (Asteraceae)

Aster ageratoides var. pendulus, a recently described taxon, is endemic to Mt Hupingshan of north‐western Hunan, China. Field observations and collections were made from the only known population. Root‐tip squashes were used to determine the chromosome numbers of 96 plants and 61 seedlings from the achenes of eight sample plants. The results show that var. pedulus is a swarm of 30 cytotypes with nearly continuous chromosome numbers from 2n = 60 to 2n = 92. Chromosome numbers of 61 seedlings vary from 2n = 61 to 2n = 91, belonging to 18 cytotypes. The chromosome number variation of var. pendulus is highly unusual not only in the A. ageratoides polyploid complex but also in angiosperms. Such an enormous continuous variation of chromosome numbers could have arisen by the combined effect of hybridization, recent origin and high levels of polyploidy. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 165 , 378–387.     

CHROMOSOME NUMBERS IN THE LEGUMINOSAE. III. SPECIES OF THE SOUTHWESTERN UNITED STATES AND MEXICO

Turner, B. L., and Olin S. Fearing. (U. Texas, Austin.) Chromosome numbers in the Leguminosae. III. Species of the Southwestern United States and Mexico. Amer. Jour. Bot. 47(7) : 603–608. Illus. 1960.—Chromosome counts for 43 species of the Leguminosae from the southwestern United States and Mexico have been reported. These include first reports for 42 taxa of which 16 are for the subfamily Mimosoideae. Olneya tesota (2n = 18) is the only new generic count listed. Chromosome reports of particular significance include a single polyploid count for a North American species of Acacia, as well as diploid and tetraploid counts for closely related taxa in this genus. Four species of the genus Schrankia were found to be diploid with In = 26, indicating a base of x = 13 instead of the x = 8 reported by some previous workers. Leucaena pulverulenta was found to have a diploid count of 2n = 56 indicating a base of x = 14.     

Conservation of chromosomal location of nucleolus organizer in American marsupials (Didelphidae)

The distribution and expression of nucleolus organizer regions (NORs) were analyzed in seven species of marsupials representative of the three karyotypes (2n = 14, 18 and 22) found in the American family Didelphidae. Analyses comprised silver-staining of NORs and fluorescence in situ hybridization with an rDNA probe. In addition to confirming the variability in number and distribution of NORs in Didelphidae, we demonstrated the conserved location of NORs on one autosome pair in the three karyotypes. In Monodelphis domestica (2n = 18), the NOR on the X chromosome was not inactivated in females.     

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.     

Chromosome number and DNA ploidy level reports from Central Europe — 3

Third part of chromosome number and DNA ploidy level reports from Central Europe comprising whole Carpatho-Pannonian region includes the data for following taxa: Scilla bifolia s. str. (2n = 18), S. bifolia agg. (2n = 36, 54), S. drunensis subsp. drunensis (2n = 36), S. drunensis subsp. buekkensis (2n = 36), S. kladnii (2n = 18) and S. vindobonensis (2n = 18) by J. Kochjarová from Austria, Slovakia and the Czech Republic (nos. 27–32); Campanula macrostachya (2n = ca 32) and Erysimum diffusum (2n = 14, 28) by E. Michalková from Hungary (nos. 33–34). Original unpublished reports should be sent to the editor on following address: patrik.mraz@upjs.sk. For further details and arrangements of reports see the first part (Mráz 2005). Previous parts of the reports were published in 2005 and 2006 (Mráz 2005, 2006).     

Karyomorphological studies of theIxeris dentata complex on Mount Ishizuchi

The karyotypes of four taxa in theIxeris dentata complex collected from Mt. Ishizuchi were investigated.I. dentata subsp.dentata var.dentata f.dentata had 2n=21 in all the examined individuals and its karyotypes were classified into four different types. Var.dentata f.amplifolia showed 2n=21 in all individuals and had three different karyotypes.I. dentata subsp.dentata var.albiflora showed 2n=21 and two different karyotypes.I. dentata subsp.alpicola showed three different karyotypes with 2n=21 and four different ones with 2n=28. Judging from the results of the karyotype analyses, the morphological variation of chromosomes in theI. dentata complex is assumed to be in the process of increasing asymmetry in the shape of chromosomes. Following Levitzky's principle, it can be said that the a-, b-, and c-chromosomes exist in different advanced stages in the process of chromosome variation. The c-group chromosomes are at the most advanced stage in morphological variation, the b-group in the second and the a-group in the least advanced stage. The above-mentioned structural diversity of chromosomes may be related to apomictic reproduction of theI. dentata complex.     

A karyological study of some Dianthus L. species (Caryophyllaceae) in Northeast of Iran

This study focuses on karyological investigation of four taxa of Dianthus (Caryophyllaceae) including D. crossopetalus, D. orientalis subsp. stenocalyx, D. orientalis subsp. gorganicus, and Dianthus sp. distributed in Northeast of Iran. The karyotype asymmetry/symmetry was evaluated using two methods: 1) CVCL (coefficient of variation of chromosome length) and CVCI (heterogeneity of the centromeric index); and 2) MCA (mean centromeric asymmetry). The karyotype asymmetry was also used to investigate the relationships among the taxa. Results obtained from the current study revealed that there are two different ploidy levels (2n = 2x = 30 and 2n = 4x = 60) among the investigated taxa. The indices CVCL and MCA described karyotype asymmetry correctly based on variation in chromosome length. Diagram of CVCL vs. MCA seems to be appropriate for karyological delimitation and taxonomic relationships among the Dianthus taxa under study. (© 2014 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Karyological investigations inRanunculus subg.Batrachium (Ranunculaceae) on the Aegean islands

This paper is the first report of an investigation on differentiation and evolutionary pattern inRanunculus subg.Batrachium mainly on the Aegean islands and treats the karyology in the subgenus. The following taxa and chromosome numbers are recorded.R. peltatus subsp.peltatus: 2n = 32, subsp.baudotii: 2n = 32, subsp.saniculifolius: 2n = 16, 32, and 40;R. peltatus s.l.: 2n = 32 and 48;R. penicillatus subsp.pseudofluitans: 2n = 16;R. rionii: 2n = 16;R. sphaerospermus: 2n = 16;R. trichophyllus: 2n = 32, 40, and 48;R. tripartitus: 2n = 48. The karyotypes show certain interspecific differences in chromosome size, in number of chromosome types according to centromere position and in occurrence and position of satellite chromosomes. r-Index and RCL have been calculated.     

CYTOGENETIC STUDIES OF SOME HYPOCHOERIS SPECIES (COMPOSITAE) FROM BRAZIL

Karyotypes of nine Brazilian taxa of genus Hypochoeris were studied utilizing root-tip mitotic metaphases. Two distinct groups were found. One group includes six species that showed high asymmetric bimodal karyotypes, while the second group has two species that have a karyotype similar to those observed in European species. All the species have karyotypes with 2n = 8 that are very uniform within each group, with only small morphological differences. Nucleolar organizing region and C-band patterns are shown for H. brasiliensis.     

GIEMSA C-BANDED KARYOTYPES OF SEVEN NORTH AMERICAN SPECIES OF ALLIUM

The karyotypes of seven North American Allium species were studied by Giemsa C-banding technique. Two species (A. shoenoprasum and A. tricoccum) were diploids with 2n = 16 chromosomes. Three species (A. cernuum, A. douglasii and A. geyeri) were also diploids but with 2n = 14 chromsomes. Diploid and tetraploid populations of A. textile (2n = 14, 28) were studied. The population of A. canadense studied here was a tetraploid (2n = 28). All these North American species, except A. geyeri, possessed centromeric bands in all their chromosomes and nucleolar constriction bands in their satellited chromosomes. Allium shoenoprasum contained telomeric bands in most of its chromosomes but the other species had them only in a small number of chromsomes. Only three species (A. shoenoprasum, A. textile and A. tricoccum) were found to have intercalary bands in some chromosomes. The heterochromatin distribution in B chromosomes of three species was also observed. In A. cernuum, the heterochromatin occupied most of the length of all its Bs, but in A. shoenoprasum, heterochromatin was concentrated in the centromeric region. One population of A. textile (CC1179) was found to have a B chromosome in which very little heterochromatin existed.     

Karyomorphology and nuclear DNA content of sixteen Ophrys L. taxa from Turkey

Karyotypes in 16 representative taxa of the Ophrys genus are compared, based on Feulgen-stained somatic metaphase chromosomes. The karyotypes of O. omegaifera subsp. israelitica, O. ulupinara, O. lycia, O. argolica subsp. lucis, O. argolica subsp. lesbis, O. climacis and O. reinholdii subsp. reinholdii are described for the first time. Karyological analyses indicate relationships among the species with respect to their asymmetry indices. All Ophrys taxa studied were diploid with 2n = 2x = 36 chromosomes. One B chromosome has been detected among the chromosomes of O. argolica subsp. lucis. All karyotypes are symmetrical, consisting of metacentric and submetacentric chromosomes. The longest chromosomes of all the investigated specimens contain a secondary constriction. It is determined that there is a correlation between the total number of chromosomes having secondary constrictions and the evolutionary development order of the taxa. Based on nuclear DNA content, analysis was carried out by flow cytometer using propodium iodide as fluorochrome, 2C nuclear DNA content of 16 Ophrys species varying between 20.80 pg (O. argolica subsp. lucis) and 23.11 pg (O. omegaifera subsp. israelitica). Karyotype asymmetry relationships are discussed according to the bidimensional scatter plots of A₁–A₂, CV_CL–CV_CI, CV_CL–M_CA and CV_CI–M_CA.     

SEGREGATION IN NEW ALLOPOLYPLOIDS OF GOSSYPIUM. V. MULTIVALENT FORMATION IN NEW WORLD X ASIATIC AND NEW WORLD X WILD AMERICAN HEXAPLOIDS

The New World tetraploid cottons, G. hirsutum and G. barbadense, are natural amphidiploids (genome formula, 2[AD]) combining species of the cultivated Asiatic (2A) and wild American (2D) groups of diploid cottons. Multivalent frequency, per cell, for 2 New World X Asiatic synthetic hexaploids, G. hirsutum X G. arboreum and G. barbadense X G. arboreum, is 6.68 and 7.80, respectively. Multivalents per cell for a series of New World X wild American synthetic hexaploids are: New World X G. harknessii, 3.65;— X G. armourianum, 3.96;— X G. aridum, 3.48;— X G. lobatum, 3.66; — X G. gossypioides, 1.13. The expected correlation between multivalent frequency and genetic segregation (e.g., high multivalent frequency = high recovery of recessives, and vice versa) for these hexaploids is realized for the near-allopolyploids (New World X wild American) but only approximated for the near-autopolyploid combinations, New World X Asiatic. This is explained on the basis that different homogenetic:heterogenetic bivalent ratios are expected in autopolyploids as compared to allopolyploids.     

Karyosystematics ofVeronica sect.Beccabunga (Scrophulariaceae) with special reference to the taxa in Turkey

The chromosome numbers ofVeronica beccabunga subsp.abscondita (2n=18),V. anagallis-aquatica subsp.lysimachioides (2n=18),V. anagallis-aquatica subsp.michauxii (2n=36), and of a presumable hybridV. poljensis × V. anagalloides (2n=36) are new for the relevant taxa. — A new number was found forV. anagalloides subsp.heureka (2n=18). The other counts, ofV. scardica (2n=18),V. anagalloides subsp.anagalloides (2n=18), andV. anagallis-aquatica subsp.oxycarpa (2n=36) agree with earlier findings.—Data from the literature have been critically evaluated, in some cases the relevant vouchers have been checked and, where needed, the original determinations have been revised. For several taxa, comments concerning their systematics and remarks on their distribution in Turkey are given.