Cytotaxonomische Untersuchungen in der Artengruppe desRanunculus alpestris (Ranunculaceae)

Morphological and cytological investigations as well as crossing experiments were carried out with the 5 species of theRanunculus alpestris group (R. alpestris L.,R. traunfellneri Hoppe,R. bilobus Bertol.,R. crenatus Waldst. etKit.,R. magellensis Ten.). A key to the species is presented; localities and distribution are given in addition to extensive diagnoses. Crossing experiments between the 5 taxa were successful (F₁-F₃ individuals, backcross types, tripeland quadrupelbastards); the morphology of the experimentally obtained F₁-hybrids was mostly intermediate. All 5 species as well as all hybrids have a chromosome number of 2n = 16; there is no statistically significant difference between the karyotypes of the 5 taxa. According to the results of the morphological investigations and the crossing experiments we can distinguish 2 subunits of very closely related species: a)R. alpestris, R. traunfellneri, R. bilobus; b)R. crenatus, R. magellensis. The speciation within the group ofRanunculus alpestris is discussed.

     

Vergleichende cytotaxonomische Untersuchungen anRanunculus seguieri und der Artengruppe desR. alpestris (Ranunculaceae)

Morphological and cytological investigations as well as crossing experiments were carried out withRanunculus seguieri Vill. and 4 species of theRanunculus alpestris L. group (R. alpestris L.,R. traunfellneri Hoppe,R. bilobus Bertol.,R. crenatus Waldst. & Kit.). ForR. seguieri andR. alpestris, localities and distribution are given in addition to extensive diagnoses. A key to the species includes morphological characteristics and distribution data forR. traunfellneri, R. bilobus, andR. crenatus. New diagnostic characters are described. Crossing experiments betweenR. seguieri and the species of theR. alpestris group were unsuccessful. All 5 species have a chromosome number of 2n = 16, the record forR. bilobus is new. There is no statistically significant difference between the karyotypes ofR. seguieri andR. alpestris s. str. Nevertheless, according to morphological evidence and crossing experiments,R. seguieri is not closely related to theR. alpestris group.

     

Experimental evidence on the affinities ofPapaver somniferum (Papaveraceae)

Results obtained from crossing experiments betweenP. somniferum subsp.somniferum (2n = 22) and subsp.setigerum (2n = 44),P. glaucum (2n = 14) andP. gracile (2n = 14) and from the observation of meiotic chromosome pairing in the various hybrids obtained do not provide straightforward evidence for the hypothesis thatP. somniferum originated as a triploid hybrid between taxa similar toP. glaucum andP. gracile (Kadereit 1986a, b).—On the one hand, the pattern of crossability found reflects the closer similarity of subsp.somniferum toP. glaucum and of subsp.setigerum toP. gracile, which was interpreted as segregation of parental characters, and the high frequency of 2n = 28 chromosomes among F₂-progeny from the hybrid subsp.somniferum × subsp.setigerum (2n = 33) might reveal n = 7 as the base number also ofP. somniferum. On the other hand, however, the general difficulty of obtaining hybrids, and the low incidence of bivalent formation in their meiosis, probably indicating a lack of chromosome homology between the different species, do not fit the above hypothesis.—These results are in marked contrast to the morphological similarity between the three species involved.     

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.     

Chromosome numbers of European blackberries (Rubus subg.Rubus,Rosaceae)

Chromosome numbers are presented for 32 collections of 29 European blackberry species (Rubus subg.Rubus) from Germany. One species is triploid (2n = 21), 27 species are tetraploid, (2n = 28), and one species is pentaploid (2n = 35). Chromosome numbers are reported for the first time ofR. adspersus, R. amisiensis, R. calvus, R. conothyrsoides, R. contractipes, R. demissus, R. elegantispinosus, R. ferocior, R. foliosus, R. hypomalacus, R. leucandrus, R. nemorosus, R. platyacanthus, R. praecox, R. rhombifolius, andR. rhytidophyllus. Chromosome numbers forR. dasyphyllus, R. gelertii, R. glandithyrsos, R. lamprocaulos, R. lindebergii, R. macrophyllus, R. montanus, R. muenteri, R. pedemontanus, R. polyanthemus, R. senticosus, R. silvaticus, andR. vigorosus are confirmed.     

Reducing the tetraploid non-nodulating alfalfa (Medicago sativa) MnNC-1008(NN) germ plasm to the diploid level

MnNC-1008(NN) (referred to as MN-1008) is a tetraploid alfalfa mutant with two recessive genes (nn ₁ and nn ₂ )conditioning the non-nodulating trait. The tetraploid level (2n=4x=32) of this Medicago sativa germ plasm was reduced to the diploid (2n=2x=16) level using the 4x-2x genetic cross originally described as a workable method for the induction of haploidy in alfalfa by T. E. Bingham. In our experiments more than 7000 emasculated flowers of a single non-nodulating MN-1008 mutant alfalfa plant with purple petals were cross-pollinated with pollen from a single, diploid, yellow-flowered alfalfa plant. Mature seeds from these crosses were collected and germinated, after which the plants were subjected to morphological and cytogenetic analyses as well as to DNA fingerprinting. Out of 26 viable progeny, 6 were hybrid plants, 19 proved to be self-mated derivatives of MN-1008, while one descendant turned out to be a diploid (2n=2x=16), purple flowered, non-nodulating plant denoted as M. sativa DN-1008. This diploid, non-nodulating alfalfa plant can serve as starting material to facilitate the comprehensive morphological, physiological and genetic analysis (gene mapping and cloning) of nodulation in order to learn more about the biology of the symbiotic root nodule development. To produce diploid, nodulating hybrid F₁ plants, DN-1008 was crossed with a diploid, yellow-flowered M. sativa ssp. quasifalcata plant. An F₂ population segregating the nn ₁ and nn ₂ genes in a diploid manner, in which the genetic analysis is more simple than in a tetraploid population, can be established by self-mating of the F₁ plants.     

Cytotaxonomy and breeding system of the genusBiscutella (Cruciferae)

Chromosome counts were determined for 46 populations ofBiscutella representing 28 taxa. The genus was found to contain diploid taxa with 2n = 12, 16 and 18, tetraploid taxa with 2n = 36 and hexaploid taxa having 2n = 54.B. laevigata L. s. l. consists of diploid and tetraploid populations which are poorly differentiated morphologically. TetraploidB. laevigata s. l. and hexaploidB. variegata Boiss. & Reuter (s. l.) are characterized by chromosomal instability. The variation in chromosome numbers and the occurrence of polyploidy is discussed in relation to the taxonomy of the genus. An investigation of the breeding system showed that most of the annual species were self-compatible and partly inbreeding and most of the perennial species self-incompatible and, therefore, outbreeding, while one annual species,B. cichoriifolia Loisel., showed both systems.     

Karyosystematic studies on threeCrepis species (Asteraceae) endemic to Greece

This work examines the cytogeographical distribution, the morphological characters, and the karyotypes of threeCrepis species endemic to Greece (C. sibthorpiana, C. incana, andC. heldreichiana). C. sibthorpiana is diploid (2n = 2x = 8),C. incana is diploid (2n = 2x = 8) and tetraploid (2n = 4x = 16, 17), andC. heldreichiana is always dekaploid (2n = 10x = 40). The Giemsa positive bands, usually pairs of dots, are mainly centromeric inC. incana, while they are terminal inC. sibthorpiana (on the short arm of all chromosomes) and inC. heldreichiana (on both arms of all chromosomes). Intercalary C-bands are scarce and usually variable within karyotypes, individuals, and species. The most variable karyotype both in Feulgen and Giemsa preparations is that ofC. incana, in which also supernumerary chromosomes were observed, which are polysomic to standard set members. On the basis of morphological and karyological data the evolutionary relationships among the threeCrepis taxa are discussed.     

Fertility relationships ofGeranium (Geraniaceae): sectt.Ruberta,Anemonifolia, Lucida andUnguiculata

Cross-pollinations were carried out among 11 briefly described species ofGeranium. Eight species pairs produced hybrids, of which five had not been reported before. The close relationship ofG. purpureum, G. robertianum andG. rubescens (sect.Ruberta) was confirmed; they form a polyploid series (diploid, tetraploid and octoploid on base x = 16). ForG. canariense (sect.Anemonifolia), another octoploid on base x = 16, the results suggest greater affinity with the former section.G. maderense andG. palmatum of sect.Anemonifolia (2n = 68) are confirmed as closely related to each other.G. maderense produced hybrids withG. robertianum (2n = 64; sect.Ruberta) and withG. cataractarum (2n = 36; sect.Unguiculata). Meiosis in the latter hybrid suggests allopolyploidy between parents with 2n = 32 and 2n = 36. Whereas all these species clearly form a very close alliance,G. lucidum (sect.Lucida) andG. macrorrhizum andG. dalmaticum (both sect.Unguiculata), appear genetically more isolated from them. One plant ofG. macrorrhizum ×G. dalmaticum was raised. — In crosses where hybrids or non-germinating seeds resulted, the reciprocal cross in the majority of cases produced a greatly inferior result or none at all. This asymmetry of response could in some cases be explained by inability of pollentubes from short-styled parents to reach the ovary of a long-styled species and in others by a modification of Hogenboom's theory of incongruity, but neither explanation works for every case. In all our asymmetric results where the ploidy level differed the diploid was the successful female, not the tetraploid, as is usually the case. — Some variation in results from year to year could be attributed to weather conditions.Dedicated to Hofrat Univ.-Prof. DrK. H. Rechinger on the occasion of his 80th birthday.     

Karyomorphological analyses and heterochromatin characteristics disclose phyletic relationships among 2n = 32 and 2n = 36 species ofOrchis (Orchidaceae)

The karyotypes of eight taxa ofOrchis L. with 2n = 32 and 2n = 36 have been investigated using morphometrical measurements following staining with Feulgen, Giemsa (C-banding) and the DNA specific fluorochrome Hoechst 33258. The karyotypes ofO. coriophora subsp.fragrans, andO. papilionacea proved to be the most asymmetrical, whileO. morio andO. longicornu exhibited the most symmetrical karyotypes. Using C-banding and the fluorochrome H33258 only the taxa with high asymmetry indices showed the presence of differentially stained chromatin bands. In most chromosomes heterochromatin bands were present at the telomeric position. The present results seem to indicate that the analysed species do not form a homogeneous group and further subdivisions are possible, which, in turn, do not always correlate with divisions based on morphological characters. Both karyomorphology and heterochromatin distribution coincide in indicating a possible evolutionary pathway.     

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.     

Stabilization of tetraploid triticale with chromosomes from Triticum aestivum (ABD)(ABD)RR (2n = 28)

Summary F₁ hybrids with the genome constitution ABDERR (2n = 6x = 42) or ABDE(AB)RR (2n = 7x = 49), selected from crosses between either an octoploid Triticum aestivum/Thinopyrum elongatun amphiploid and tetraploid Secale cereale (AABBDDEE x RRRR) or autoallohexaploid triticale [AABBDDEE x (AB)(AB)RRRR], were backcrossed to tetraploid triticale (AB)(AB)RR and selfed for six generations. Thirty-three different tetraploid F₆ progenies were karyotyped using C-banding. The aneuploidy frequency was 6.6% with 4.0% hypoploids and 2.6% hyperploids. Among 71 plants with 28 chromosomes, 53.5% had a stabilized karyotype while 46.5% were unstabilized with at least one homoeologous group segregating for A-, B-, or D-genome chromosomes. The stabilized plants represent 19 different tetraploid karyotypes with six of them not containing any detectable D-genome chromosomes from T. aestivum or E-genome chromosome from Th. elongatum. Thirteen lines were (ABD)(ABD)RR tetraploids with one-to-three disomic substitutions of D-genome chromosomes for A or B-genome chromosomes. No disomic substitution of E-genome chromosomes was identified. On average 0.58 D substitutions per line were determined. Of the seven D-genome chromosomes only four, 1D, 2D, 5D, and 7D, were present in their disomic state. In unstabilized karyotypes, chromosomes 3D, 4D, and 6D were present in their monosomic state. Among all 30 viable plants (42.3%), the order of decreasing frequency of Dgenome chromosomes was 5D (25.0%), 1D (20.0%), 2D (10.0%), 6D (5.0%), and 3D (1.7%). Plants with 4D and 7D chromosomes were not viable. An increase in the number of D-genome chromosomes in the (ABD) genome is associated with a decrease in viability and fertility. Minor differences in the C-banding of chromosomes in homoeologous groups 1, 5, and 6 indicate the possibility of translocations between A-, B-, D-, and E-genome chromosomes. Evolutionary and breeding aspects of tetraploid triticale with mixed genomes are discussed.     

Chromosome numbers and taxonomic implications in the fern genusAzolla (Azollaceae)

Investigation of chromosome numbers of allAzolla species, and for the first time of hybrids, has been undertaken. Removal of wax from the leaf surface proved invaluable in achieving clear cytological preparations and providing unambiguous chromosome numbers. In contrast to previous records, the speciesA. pinnata, A. filiculoides, A. filiculoides var.rubra, A. caroliniana, A. microphylla, andA. mexicana were found to be 2n=44, andA. nilotica to be 2n=52. Several triploids (2n=66) and one tetraploid (2n=88) were identified. No geographical pattern could be observed in the distribution of triploids which probably derive from the function of unreduced gametes. The chromosome number of hybrids occasionally deviates from the diploid chromosome number (2n=44). The small chromosome size limits karyotypic analysis and only differences in overall chromosome size can be observed. Taxonomic implications of chromosome numbers and sizes are discussed.     

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.     

Giemsa karyotypes and their evolutionary significance inScilla bifolia,S. drunensis,andS. vindobonensis (Liliaceae)

Quantitatively evaluated C-banded and conventional karyograms are presented for the related speciesScilla bifolia subsp.danubialis Speta,S. drunensis (Speta)Speta, andS. vindobonensis Speta. On the basis of banding patterns and karyotype structureS. bifolia subsp.danubialis (2n = 18, 2×) andS. drunensis (2n = 36, 4×) are quite similar, whileS. vindobonensis (2n = 18, 2×) is entirely different. There is a moderate degree of karyotypic variation withinS. bifolia subsp.danubialis andS. drunensis. However, inS. vindobonensis karyotypes and banding patterns are almost completely stable over a geographical range of about 500 km. The present results confirm the recent taxonomic separation ofS. vindobonensis fromS. bifolia, and suggest a considerable phylogenetic distance between these two diploid species. The results are discussed with reference to the morphological characters of the species and their geographical distribution.Evolution ofScilla and Related Genera, II.Dedicated to Univ.-Prof. Dr.Elisabeth Tschermak-Woess on the occasion of her 60th birthday.     

Interspecific hybridizations among species of theElymus semicostatus andElymus tibeticus groups (Poaceae)

Interspecific hybridizations were made between species of theE. semicostatus group, viz.,E. semicostatus (Nees exSteud.)Meld.,E. validus (Meld.)B. Salomon,E. abolinii (Drob.)Tzvel., andE. fedtschenkoi Tzvel., and species of theE. tibeticus group, viz.,E. pendulinus (Nevski)Tzvel.,E. tibeticus (Meld.)Singh,E. shandongensis B. Salomon, andE. gmelinii (Ledeb.)Tzvel., as well as among species within theE. tibeticus group. All species are tetraploid (2n = 4x = 28) and possess SY genomes. Meiotic pairing data from 24 hybrids involving 17 interspecific combinations are presented. The average number of chiasmata per cell ranged from 17.91 to 26.20 in hybrids within theE. tibeticus group, compared with 7.26 to 22.04 in hybrids between the two species groups. Despite the extensive collection of cytological data, there was no definite evidence for confirming or disproving the separate existence of the two groups.     

C-banding variation in the Moroccan oat species Avena agadiriana (2n=4x=28)

The C-banding technique was used to describe the chromosomes of a relatively recently-discovered Moroccan oat species, Avena agadiriana (2n=4x=28). A substantial amount of polymorphism for arm ratios and C-banding patterns was observed among five accessions of this species. However a common set of ten putatively homologous chromosomes was identifiable among the five accessions. The chromosomes of A. Agadiriana do not closely match those of any of the previously described diploid or tetraploid oat species in terms of their arm ratios and C-banding patterns. However, their overall C-banded appearance generally resembles the A/B/D groups of chromosomes of Avena species, rather than the more hetrochromatic C genomes. Implications of these findings in terms of chromosome evolution in the genus Avena are discussed.Contribution no. 95-490-J of the Kansas Agricultural Experiment Station, Kansas State University, Manhattan, KS, USA     

Drei neue Arten desFestuca ovina-Formenkreises (Poaceae) aus dem Osten Österreichs

WithinFestuca ovina agg. two hexaploid taxa:F. brevipila andF. carnuntina, and one tetraploid:F. eggleri, are described as new species.F. pseudovina Hack. exWiesb. is better treated asF. valesiaca Schleich. exGaudin subsp.parviflora (Hack.), comb. nova.

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Drei neue Arten desFestuca ovina-Formenkreises (Poaceae) aus dem Osten Österreichs

     

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.     

Ranunculus cacuminis andR. crenatus,representatives of theR. alpestris-group (Ranunculaceae) on the Balkan Peninsula

Morphological and karyological investigations as well as crossing experiments were carried out withR. cacuminis, R. crenatus, andR. alpestris. A key is presented including all 6 taxa of theR. alpestris-group. Ecological and geographical data for the Balkan Peninsula are also presented.R. cacuminis is diploid with 2n = 16 chromosomes as all taxa of the group, its karyotype exhibits no statistically significant differences to those from the 5 other taxa. In crossing experiments betweenR. cacuminis andR. alpestris, no hybrids could be obtained, whereas those betweenR. cacuminis andR. crenatus were slightly successful. According to its morphology, geographical distribution, and crossing behaviour,R. cacuminis is closely related toR. crenatus, and probably has originated fromR. crenatus by quantum speciation.