Cytogenetics, geographical distribution, and pollen fertility of diploid and tetraploid cytotypes of Santolina pectinata Lag. (Asteraceae: Anthemideae)

A cytogenetic study of 62 populations of Santolina pectinata in Spain shows the existence of two ploidy levels. The diploid cytotypes with 2 n  = 18 occupy the eastern Betic mountains, and the tetraploid cytotypes with 2 n  = 36 are located on the spurs of the Iberian System. The former show a much wider ecological spectrum than the latter. Mixed cytotypes were observed in two diploid populations, with one tetraploid in each, showing different karyotypes. Three trisomic individuals were detected, one in a diploid population and the other two in a tetraploid population. Also, three hypotetraploid individuals were detected in a tetraploid population. Polyploidy is shown to be spontaneous and recurrent, promoting partial sterility in the pollen. Structural chromosomal changes, principally translocations, and local speciation through autopolyploidy are the principal factors in the evolution and diversification of this species.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 657–667.     

A review of ploidy in the genus Prosopis (Leguminosae)

The genus Prosopis contains 44 species of trees and shrubs, the majority of which originate in the Americas. Most species are reported to be diploid, with a somatic chromosome number of 2 n  = 28. There are rare reports of polyploidy, although it is thought that these may represent polysomaty in root tissues. However, flow cytometry has recently indicated that P. juliflora is entirely tetraploid with a somatic chromosome number of 2 n  = 56. In order to clarify the situation, an extensive review of ploidy in Prosopis was undertaken, the first of its kind. The ploidy levels of 124 samples of Prosopis from 21 countries, including both the natural and introduced ranges, were analysed using flow cytometry. In addition, a comprehensive literature review was carried out, examining 305 published ploidy values and covering 32 of the 44 species of Prosopis . Flow cytometry analysis suggests that P. juliflora is the only tetraploid species, with a somatic chromosome number of 2 n  = 4 x  = 56, whilst the remainder of the species analysed are diploid with 2 n  = 2 x  = 28, including the first report for P. articulata (2 n  = 28). A critical review of published ploidy values shows that all species of Prosopis are reported to be entirely diploid, except P. glandulosa , P. juliflora , and P. koelziana , for which both diploid and tetraploid values have been recorded. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 425–438.     

Ploidal levels in the arctic-alpine polyploid Draba lactea (Brassicaceae) and its low-ploid relatives

Ploidal level information is of particular importance in intricate polyploid complexes such as in arctic-alpine Draba . Relative DNA content is reported for the tetra- and hexaploid D. lactea and seven of its low-ploid relatives. Flow cytometry was used to study 200 plants from 93 populations, the screening based on relative fluorescence. Absolute DNA content was determined by Feulgen densitometry for 13 plants from seven species, and reference chromosome numbers were determined in 12 plants (1–3 per species) representing six species. The plants grouped into diploids (2 n  = 16), tetraploids (2 n  = 32), hexaploids (2 n  = 48), and two triploids. Each ploidal level showed a linear increase in relative DNA content, pointing to a relatively recent polyploid origin. The diploid level was confirmed in D. nivalis, D. subcapitata, D. fladnizensis , and D. lonchocarpa. Draba palanderiana , reported previously as di-, tetra- and octoploid, was diploid in all investigated accessions. Hexa- and tetraploids were observed in D. lactea , in approximately the same ratio (8 : 1) as reported previously. The ploidal levels of the Central Asian D. altaica and D. turczaninovii are reported here for the first time as diploid and tetraploid, respectively.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 333–347.     

Asynchronous meiotic rhythm as the cause of selective chromosome elimination in an interspecific <Emphasis Type="Italic">Brachiaria</Emphasis> hybrid

This paper reports the occurrence of chromosome elimination during microsporogenesis in an interspecific hybrid between a sexual diploid accession (SEX) of Brachiaria ruziziensis (2n=2x=18) and an apomictic tetraploid accession (APO) of B. brizantha (2n=4x=36). Meiosis was very abnormal in the triploid hybrid (2n=3x=27); we observed a distinct asynchrony from metaphase I to the end of meiosis. The APO and the SEX genomes did not show the same meiotic rhythm. When the former, with nine bivalents, was in metaphase I, the nine SEX univalents were not yet aligned; when the latter reached the plate, the APO genome was already in anaphase. In subsequent stages, the APO genome had reached the poles while the SEX was undergoing sister-chromatid segregation. As the SEX genome always remained temporally behind, it gave rise to one extra-nucleus in each pole. In the second division, the behavior was the same but anaphase II did not occur for the SEX genome, and only one extra-nucleus was observed in each cell in telophase II. Chromosome elimination for the SEX genome ranged from partial to total. The importance of these findings with respect to Brachiaria breeding programmes is discussed.     

Polyploidy in Aconitum subgenus Lycoctonum (Ranunculaceae)

Chromosome numbers and morphology in 22 populations belonging to 11 taxa of Aconitum subgenus Lycoctonum (Ranunculaceae) from China were studied. Some taxa were diploid, with 2 n  = 16, but four species and two varieties were found to be tetraploid, with 2 n  = 32. They are concentrated in the Hengduan Mountains region in south-west China, indicating that polyploidy could have played an important role in the speciation of the subgenus in this region, one of the areas of the world with a high concentration of endemic species. The relationships of some of the species are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 343–353.     

Karyological and taxonomic notes on Ophrys (Orchidoideae, Orchidaceae) from the Iberian Peninsula

Karyological information on Iberian Ophrys species is very limited. This paper provides the haploid and diploid chromosome numbers of 11 taxa of sect. Pseudophrys and sect. Ophrys , both of which are well represented in the Iberian Peninsula, and two taxa from Tunisia. The first data on chromosome numbers for O. vasconica (2 n  = 72, 74), O. ficalhoana (2 n  = 36), O. picta (2 n  = 36), O. sphegifera ( n  = 18, 2 n  = 36, 38) and O. passionis (2 n  = 36) are also presented, confirming the stability of the chromosome number in Ophrys . In addition, populations of the group O. omegaifera ( O. dyris and O. vasconica ), together with tetraploidy, pentaploidy and the existence of aneuploid phenomena, are reported for the first time in Iberia. The basic diploid number is always 2 n  = 36. The karyotypes of several species were analysed. Evolutionary trends in Ophrys chromosomes are discussed. Taxonomic and phytogeographical details are provided on several species or groups of species from Iberia.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003 , 142 , 395−406.     

Karyotype analysis in South American species of Myrtaceae

In Myrtaceae (Myrteae), the diploid chromosome number 2 n  = 2 x  = 22 is the most common, although variations of ploidy level occur, with some triploid (2 n  = 3 x  = 33) and tetraploid (2 n  = 4 x  = 44) records. Karyotype details in this group are scarce because the chromosomes are small (< 2 μm). In this work, we carried out a karyotypic analysis of 15 species of Myrtaceae grouped in different subtribes and genera. Measurements of chromosome length (long arm, L ; short arm, S ) were taken and several karyotypic parameters were calculated for each species. The karyotypes in fleshy-fruited taxa (Myrteae) were more varied than in the other previously analysed dry-fruited group ( Eucalyptus , Eucalypteae), in which the chromosomes were exclusively metacentric.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155 , 571–580.     

Karyological observations on plant taxa endemic to the Balearic Islands

Mitotic chromosome numbers are reported from 35 vascular plant taxa endemic to the Balearic Islands that are poorly known cytogenetically. The chromosome numbers of Chaenorhinum rodriguezii (2 n  = 14), Coristospermum huteri (2 n  = 22), Carduus ibicensis (2 n  = 26), Filago petro-ianii (2 n  = 26), Ononis crispa ssp. zschackei (2 n  = 30), and Ophrys bertolonii ssp. balearica (2 n  = 36) were determined for the first time. New chromosome numbers are reported in Pimpinella bicknellii (2 n  = 22), Calamintha rouyana (2 n  = 46), Vicia bifoliolata (2 n  = 14), and Teucrium asiaticum (2 n  = 30). A new diploid cytotype (2 n  = 16) was found in Ranunculus paludosus ssp. barceloi , which is restricted to populations from the western Balearics (Eivissa and Formentera), whereas tetraploid plants are known from the eastern Balearics (Mallorca). It is suggested that the tetraploid cytotype arose from the diploid cytotype through autopolyploidy. The idiograms of Coristospermum huteri , Pastinaca lucida , Pimpinella bicknellii , Carduus ibicensis , Helianthemum scopulicolum , Helichrysum crassifolium , Vicia bifoliolata , p aeonia cambessedesii , Helleborus lividus , Ranunculus paludosus ssp. barceloi , and Linaria aeruginea ssp. pruinosa are determined for the first time.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 463–476.     

Hybrid origins of Ixeris nakazonei (Asteraceae, Lactuceae) in the Ryukyu Archipelago, Japan: evidence from molecular data

The proposed hybrid origin of Ixeris nakazonei collected in the Ryukyu Archipelago (Okinawajima) and the Atsumi Peninsula (Koijigahama) of central Japan was examined using nuclear DNA (nDNA) and chloroplast DNA (cpDNA) markers. The results indicated that most tetraploids of I. nakazonei (2 n  = 32) had nDNA of both putative parents, I. debilis (2 n  = 48) and I. repens (2 n  = 16). The first tetraploid of I. nakazonei must have arisen directly from interspecific hybridization between I. debilis and I. repens , considering the intermediacy of the chromosome number. All tetraploids on Okinawajima had cpDNA of I. repens , while those in the Atsumi Peninsula had cpDNA of I. debilis , indicating that the tetraploid has at least two independent origins. All hexaploids of I. nakazonei (2 n  = 48), on the other hand, had cpDNA of I. debilis , but some of the hexaploids had nDNA of both putative parents. Ixeris debilis and I. repens may also be involved in the origin of the hexaploid, although the establishment process of this cytotype is still obscure. Since the hexaploid of I. nakazonei is cytologically indistinguishable from I. debilis , it can backcross with I. debilis . Introgression from diploid I. repens to hexaploid I. debilis mediated by I. nakazonei possibly occurs on Okinawajima.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 379–387.     

Determination of ploidy and nuclear DNA content in populations of Atriplex halimus (Chenopodiaceae)

Nuclear DNA contents were determined by flow cytometry for 20 populations of the perennial C₄ shrub Atriplex halimus L. (Chenopodiaceae) originating from the Mediterranean basin and Fuerteventura (Canary Islands). Two populations were also analysed for chromosome number: one (from Ibiza, Spain), with a 2C nuclear DNA content of 2.40 pg, was shown to be diploid (2 n  = 2 x  = 18), whilst the other (from Sicily, Italy), with 5.11 pg, was tetraploid (2 n  = 2 x  = 36). With respect to nuclear DNA content, two groups of populations were detected, diploids with 2.40–2.44 pg and tetraploids with 4.77–5.13 pg. The diploid populations were mainly from the western Mediterranean (Spain and France) and Fuerteventura, whereas tetraploids were generally, but not exclusively, from more arid areas in North Africa and the eastern Mediterranean. In general, the diploid and tetraploid populations corresponded to the subspecies halimus and schweinfurthii , respectively. For certain populations having morphologies intermediate between those considered typical of these two subspecies, nuclear DNA contents showed them to be tetraploid. There was significant variation in nuclear DNA content among the tetraploid populations, with greater values in the more easterly populations.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 147 , 441−448.     

Polyploidy and karyotype variation in Turkish Bellevalia (Hyacinthaceae)

There are 20 Bellevalia species in Turkey, half of which are endemic. Chromosome numbers are known for 15 species. A chromosome survey of 145 Bellevalia individuals showed that the karyotype is remarkably stable. All are based on x  = 4. The majority are diploid with 2 n  = 8, but there is also a polyploid series of 2 n  = 16, 24 and 32. Aneuploidy occurs only at the octoploid level. Eleven individuals had metacentric B chromosomes, one had acrocentric Bs and one had telocentric Bs. Bellevalia pycnantha and B. paradoxa are morphologically similar, with B. pycnantha reduced to a synonom of B. paradoxa .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 87–98.     

Meiotic behaviour in three interspecific three-way hybrids between Brachiaria ruziziensis and B. brizantha (Poaceae: Paniceae)

The meiotic behaviour of three three-way interspecific promising hybrids (H17, H27, and H34) was evaluated. These hybrids resulted from the crosses between B. ruziziensis X B. brizantha and crossed to another B. brizantha. Two half-sib hybrids (H27 and H34) presented an aneuploid chromosome number (2n = 4x = 33), whereas hybrid H17 was a tetraploid (2n = 4x = 36), as expected. Chromosome paired predominantly as multivalents suggesting that genetic recombination and introgression of specific target genes from B. brizantha into B. ruziziensis can be expected. Arrangement of parental genomes in distinct metaphase plates was observed in H27 and H34, which have different male genitors. Hybrids H17 and H34 have the same male genitor, but did not display this abnormality. In H17, abnormalities were more frequent from anaphase II, when many laggard chromosomes appeared, suggesting that each genome presented a different genetic control for meiotic phase timing. Despite the phylogenetic proximity among these two species, these three hybrids presented a high frequency of meiotic abnormalities, mainly those related to irregular chromosome segregation typical of polyploids, H34, 69.1%; H27, 56.1% and H17, 44.9%. From the accumulated results obtained through cytological studies in Brachiaria hybrids, it is evident that cytogenetical analysis is of prime importance in determining which genotypes can continue in the process of cultivar development and which can be successfully used in the breeding. Hybrids with high frequency of meiotic abnormalities can seriously compromise seed production, a key trait in assuring adoption of a new apomictic cultivar of Brachiaria for pasture formation.     

Cytogenetics of species of Chamaecrista (Leguminosae – Caesalpinioideae) native to southern Brazil

Chromosome numbers, karyotypes, meiotic behaviour and pollen analysis are presented for species of Chamaecrista Moench (Leguminosae, Caesalpinioideae, Cassieae ) native to southern Brazil: C. nictitans ssp. patellaria , C. nictitans ssp. disadena , C. repens , C. rotundifolia , C. flexuosa , C. vestita and C. desvauxii . Meiotic behaviour is reported for the first time for all the taxa and was very regular; only bivalents were formed at diakinesis and metaphase I, chromosome disjunction and segregation were regular at anaphases I and II, meiotic indexes were over 99% and pollen fertility was over 92%. Pollen grains were subprolate in C. flexuosa and C. vestita and prolate–spheroidal in the other taxa. Karyotypes were symmetrical in all six species and the data are original, except for C. nictitans ssp. p atellaria. Chromosome number is presented for the first time for C. repens (2 n  = 16) and has been confirmed for the other taxa: 2 n  = 14 for C. desvauxii , 2 n  = 32 for the tetraploid C. nictitans ssp. patellaria and C. nictitans ssp. disadena , and 2 n  = 16 for the other species. These two basic numbers found in the genus, x  = 7 and x  = 8, point to chromosome evolution by dysploidy, which has also been accompanied by polyploidy.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 429–439.     

Cytological studies on the Sino-Himalayan endemic Solms-laubachia (Brassicaceae) and two related genera

The karyotypes of six species of Solms-laubachia and one each of Parrya and Desideria (Brassicaceae) from the Himalayan-Hengduan Mountains (Tibet, Sichuan and Yunnan provinces) were investigated. Chromosome counts of S. xerophyta , S. lanata, S. platycarpa and D. baiogoinensis are reported for the first time and found to be diploid with 2 n  = 14. Diploid counts were also made of S. retropilosa , a species previously known to be a tetraploid. The karyotype formulae of D. baiogoinensis , S. eurycarpa , S. pulcherrima , S. retropilosa , S. xerophyta and S. platycarpa are 2 n  = 14 = 6 m + 6sm + 2st, whereas those of S. lanata and P. nudicaulis are 2 n  = 14 = 6 m + 2sm + 6st and 2 n  = 14 = 8 m + 4sm + 2st, respectively. Karyotypes and counts of S. eurycarpa and S. pulcherrima are in agreement with prior studies. The relationships within Solms-laubachia and among the three genera are discussed. Speciation within Solms-laubachia may have been the result of primarily intrachromosomal variation at the diploid level, and the genus probably evolved within the Himalayan-Hengduan Mountains instead of migrating from other regions.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 77−86.     

Whole-genome duplications in South American desert rodents (Octodontidae)

The discovery of tetraploidy in the red viscacha rat, Tympanoctomys barrerae (4 n  = 102) has emphasized the evolutionary role of genome duplication in mammals. The tetraploid status of this species is corroborated here by in situ PCR and Southern blot analysis of a single-copy gene. The species meiotic configuration strongly suggests a hybrid derivation. To investigate the origin of T. barrerae further, the recently described Pipanacoctomys aureus was studied. This 92-chromosome species also has a duplicated genome size, redundant gene copy number and diploid-like meiotic pairing, consistent with an event of allotetraploidization. Phylogenetic analysis of mitochondrial sequences indicates sister-group relationships between these two tetraploid rodents. The new karyotypic data and the phylogenetic relationships suggest the participation of the ancestral lineages of Octomys mimax in the genesis of P. aureus . The high overall DNA similarity and shared band homology revealed by genomic Southern hybridization as well as matching chromosome numbers between O. mimax and the descendant tetraploid species support the notion of introgressive hybridization between these taxa.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 443–451.     

Origins, establishment and evolution of new polyploid species: Senecio cambrensis and S. eboracensis in the British Isles

Two new polyploid species of Senecio have originated in the British Isles in recent times following hybridization between native S. vulgaris (2 n  = 40) and introduced S. squalidus (2 n  = 20). One of these is the allohexaploid S. cambrensis (2 n  = 60), the other is the recombinant tetraploid S. eboracensis (2 n  = 40). We review what is known about when and how each species originated, and their reproductive isolation from parents due to high selfing rates. We also review evidence that suggests S. cambrensis may have undergone rapid genome evolution since its origin, and comment on the risks of extinction to each species due to chance factors operating during the early establishment phase. The discovery of both species soon after their origin provides an unparalleled opportunity to examine two different but related forms of speciation following hybridization between the same parent species. Further detailed study of the ecology and genomics of S. cambrensis and S. eboracensis will help improve our understanding of the process of polyploid speciation in plants.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 467–474.     

Karyotype analysis in Brachiaria (Poaceae) species

This is the first karyotype characterization of Brachiaria species. Twelve accessions belonging to five species were analysed. The basic chromosome number was x = 9 and 7, the same reported for the tribe Paniceae. Variations in the chromosome number were observed in B. decumbens (2n = 18; 36) and B. humidicola (2n = 36; 42; 54). Chromosome numbers of 2n = 18 in B. ruziziensis and 2n = 36 in B. brizantha and B. jubata were recorded. Inter- and intraspecific karyotype differentiation of the accessions analysed was facilitated by variations in karyotypic symmetry. The karyotypes were generally considered symmetrical, with a tendency to asymmetry in the direction of the polyploids. It is suggested that addition, deletions and mainly polyploidy have been the most direct causes involved in the chromosome evolution of this genus.     

Ploidy variation in Buddleja L. (Buddlejaceae) in the Sino-Himalayan region and its biogeographical implications

The chromosome numbers of 27 populations of Buddleja , comprising 14 species, were counted. The basic chromosome number of all species was x  = 19, confirming previous reports. Different ploidy levels (2 n  = 38, 76, 114, 228) were observed in these taxa, representing diploids, tetraploids, hexaploids, and dodecaploids, respectively. The chromosome numbers of B. yunnanensis , B. brachystachya , and B. macrostachya are reported for the first time. The tetraploid 2 n  = 76 is a new ploidy level for B. myriantha . Particular attention was given to B. macrostachya , because of the variation in morphology and ploidy level between isolated populations of this species. Two types of interphase nuclei were recognized: the complex chromocentre type in B. macrostachya and the simple chromocentre type in the other species. Biogeographically, most of the polyploidy in the Asiatic species occurs in the Sino-Himalayan region. It seems to be associated with the uplift of the Himalayan Mountains, the orogeny of this region playing an important role in the evolution of polyploidy in these taxa.  © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 305–312.     

Chromosome numbers and karyotypes of South American species and populations of Hypochaeris (Asteraceae)

One hundred and thirty-seven new chromosome counts are reported from 104 populations of 26 native South American taxa of Hypochaeris (Asteraceae, Lactuceae), together with two invasive Mediterranean species: H. glabra and H. radicata . First reports are provided for seven taxa ( H. alba , H. cf. eremophila , H. caespitosa , H. hookeri , H. parodii , H. patagonica and H. pinnatifida ) and one new ploidy level is reported (diploid for H. incana , so far known only as a tetraploid). Including the results of this study, the chromosomes of 39 of the c . 50 Hypochaeris species known from the New World have now been counted. Most species are diploid with 2 n  = 2 x  = 8 and have bimodal, asymmetrical karyotypes. Tetraploidy (2 n  = 4 x  = 16) is reported here for the first time in H. caespitosa . Infra-specific polyploidy (probably autopolyploidy) is reported in H. incana and H. taraxacoides , both cases including infra-populational cytotype mixtures (2 x and 4 x ). Polyploidy is now known from eight South American Hypochaeris species ( c . 16%). Basic karyotype analyses allow the placement of the newly counted taxa into previously proposed but slightly modified groupings and provide the framework for further molecular cytogenetic analyses. The reported findings suggest that chromosomal change in South American Hypochaeris , in contrast to Old World species, has not involved aneuploidy, but polyploidy and/or more subtle changes in chromosome length, perhaps via satellite DNA amplification/deletion or activity of retroelements, and rDNA reorganization.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 49–60.