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.     

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.     

Chromosomal variability in the genus Primula (Primulaceae)

Chromosome numbers and measurements were recorded in 47 individuals of ten taxa of Primula , representing eight species and two subspecies from ten populations. The basic numbers of chromosomes were x  = 8, 9 or 11, and they were mostly metacentric, medium-long to medium-small, ranging in length from c . 3.6 µm to 1.6 µm. Diploid chromosome numbers of 2 n  = 2 x  = 24 and 22 were scored for the first time in P. loeseneri Kitag. and P. prenantha Balf. & Sm., respectively. A new ploidy level of 2 n  = 4 x  = 44 was found in P. burmanica Balf. & King. The recorded diploid numbers of the remaining species confirmed earlier reports. Interspecific variability in chromosome numbers was correlated with heterogeneity in their mean length. Comparison of the data with those in the literature revealed that the observed variability of chromosomal characters was compatible with the other taxonomic criteria and supports the current taxonomic delimitation. Chromosomal variation at the diploid level is the predominant feature in Primula evolution, and allopatric speciation has played a major role in its specific diversity. Subgen. Aleuritia could represent the main clade from which the other subgenera have evolved. The origin of the chromosome numbers, geographical distribution and evolution of the species were assessed, and the parallel polyphyletic mode of evolution in the genus was confirmed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 211–219.     

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.     

Chromosome numbers in Brazilian species of Crotalaria (Leguminosae, Papilionoideae) and their taxonomic significance

Chromosome numbers were counted for 23 species of Crotalaria native to Brazil. Among these data there were new counts for 15 taxa, and some confirmed previous reports or represented numbers that were different from those cited previously. The chromosome numbers most frequently found were 2 n  = 16 and 2 n  = 32. Only C. incana L. had 2 n  = 14 and C. tweediana Benth. had 2 n  = 54. The counts 2 n  = 32 and 54 were found in species of section Calycinae and 2 n  = 16 and 14 in species of section Chrysocalycinae . The data revealed the importance of chromosomal parameters in the characterization of sections Calycinae and Chrysocalycinae in Brazil. We discuss the systematic significance and evolutionary aspects for the genus, comparing the results with the two sections that are native in Brazil.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 271–277.     

Karyology of some Brazilian species of Alismataceae

The family Alismataceae ( sensu Cronquist) comprises 11 genera and 75 species of aquatic and semiaquatic herbs. In Brazil, only the two most speciose genera, Echinodorus Rich. ex Engelm. and Sagittaria L., are present. The chromosome number 2 n  = 22 is common to all species. We present karyotypes for eight species, together with their asymmetry index (TF%) and total chromatin length (TCL). All karyotypes have high karyotypic asymmetry and TCL, due to their large and mostly acrocentric chromosomes. The chromosomal evolution of the Alismatidae group is discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 159−164.     

Karyotype evolution in the genus Boronia (Rutaceae)

New chromosome counts are reported for Boronia clavata 2 n  = 14, B. heterophylla 'Near White' 2 n  = 15, B.  'Carousel' 2 n  = 16, B. deanei 2 n  = 22, B. chartacea 2 n  = 32, B. keysii 2 n  = 32, B. pilosa 2 n  = 44, B. anethifolia 2 n  = 36 and B. citriodora 2 n  = 108. Studies in 20 genotypes of 18 species and one interspecific hybrid revealed that they are highly complex in terms of chromosome number, ploidy level, chromosomal length, karyotype constitution and asymmetry. Karyotype analysis indicated that Boronia taxa with high chromosome numbers are primitive and those with lower numbers are derived. The basic chromosome number for this genus is suggested to be x = 18. Analysis of chromosome number, variations of total chromosome length (TCL) and average chromosome length (ACL), Nombre Fondamental (NF) and karyotype asymmetry suggest that dysploid reduction is the major mechanism in Boronia karyotype evolution. Chromosomal rearrangements might also have been involved. Origin, chromosome number changes and spread of Boronia are discussed in relation to the species divergence and the geological and climatic changes of the Australian continent.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 309–320.     

Cytotypes of Andropogon gerardii Vitman (Poaceae): fertility and reproduction of aneuploids

In many North American prairies, populations of Andropogon gerardii Vitman (Poaceae) are composed of hexaploid and enneaploid cytotypes (2 n  = 60, 90), with intermediates occurring occasionally. Under controlled pollination, the two common cytotypes can be crossed, producing progeny with a range of chromosome numbers. In an investigation of fertility and compatibilities of intermediate cytotypes, individuals with chromosome numbers between 60 and 90 were crossed with each other, with the 2 n  = 60 and 90 cytotypes, and with South American Andropogon species having 60 chromosomes. Regardless of cytotype, all A. gerardii plants had some fertility and virtually all crosses produced seeds. Cytotype is only partially predictive of fertility. Inter-specific hybrids between A. gerardii and South American hexaploid species were vigorous but sterile. Gene flow in natural A. gerardii populations of mixed cytotype probably involves plants of all cytotypes.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 95–103.     

The cytology of Actinidia, Saurauia and Clematoclethra (Actinidiaceae)

Meiosis and mitosis of six Chinese Actinidiaceae were studied: Saurauia tristyla DC., S. miniata C. F. Liang & Y. S. Wang, Actinidia chinensis Plach., A. deliciosa (Cheval.) C. F. Liang & A. R. Ferguson, A. indochinensis Merr. and Clematoclethra lasioclada Maxim. The chromosome numbers of Saurauia tristyla and S. miniata were 2 n  = 6 x  = 78, establishing a base chromosome number of x  = 13 in the genus, differing from the previous report of x  = 15. The chromosome number of Clematoclethra was first reported to be 2 n  = 4 x  = 48 ( x  = 12), while that of Actinidia was x  = 29, consistent with previous reports. The base chromosome number of Clematoclethra ( x  = 12) was derived from an aneuploid decrease from Saurauia ( x  = 13). Actinidia (x  = 29) was derived from the palaeotetraploid ( x  = 14), which was formed through the increase of the basic chromosome number x  = 13 to x  = 14 by aneuploidy and through the breakage of a centromere to add one more new chromosome. The chromosome data in Actinidia were consistent with the geographical and morphological evidence for the evolution of the three genera. The tropical American and Asian disjunct distribution pattern and the diversity of base chromosome numbers of Saurauia further support the probability that the genus was an early divergent from a common ancestor of Actinidia and Clematoclethra .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 369–374.     

New chromosome counts in the genus Cousinia (Asteraceae) from Iran

Chromosome counts are reported of 38 populations representing 24 species of the genus Cousinia from Iran, one of the main centres of speciation of the genus. Sixteen of the counts are new to science. Our results confirm that Cousinia exhibits a complex dysploid series ranging from x  = 13 to x  = 9. One of the basic chromosome numbers, x  = 9, is new for the genus and must be confirmed. Some considerations on the correlation between chromosome numbers and section classification are made.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 411–419.     

Reduction of chromosome number in Eleocharis subarticulata (Cyperaceae) by multiple translocations

Eleocharis subarticulata is recorded as the third species of Cyperaceae with a reduced chromosome number ( n  = 3), following reports on Rhynchospora tenuis ( n  = 2) and Fimbristylis umbellaris ( n  = 3). For Eleocharis, the numbers recorded to date vary from 2 n  = 10 to 2 n  = c. 196, with x  = 5 as the possible basic number. The karyotype of E. subarticulata was studied using conventional staining (mitosis and meiosis), C-CMA₃/DAPI banding, and FISH with 45S rDNA and telomere probes. The chromosomes showed no primary constrictions, as expected in the holocentric chromosomes of Cyperaceae. The meiotic behaviour was abnormal, with a single multivalent ring of six chromosomes at metaphase I, resulting from multiple translocations. At anaphase I six chromatids migrated to each pole, evidencing the inverted meiosis, and these groups were also visible at metaphase II. The C-CMA₃/DAPI banding technique showed only four terminal GC-rich blocks. FISH with 45S rDNA probes revealed four terminal signals, probably associated with GC-rich blocks. The telomeric probe located terminal signals in all the chromosomes, besides a hybridization site in the middle of the large pair. The occurrence of ectopic telomeric sites has not been described previously for plants with holokinetic karyotypes and with reduced chromosome numbers. These data reinforce the hypothesis of the reduction in chromosome number by multiple translocations.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 457–464.     

Cytotaxonomy of Begonia rubropunctata and B. purpureofolia (Begoniaceae)

The chromosome numbers of four Chinese Begonia species were counted for the first time as follows: 2 n  = 22 in B. rubropunctata , 2 n  = 18 in B. purpureofolia , 2 n  = 44 in B. pedatifida , and 2 n  = 22 in B. villifolia . The first two species had relatively large chromosomes with early condensed chromatin in the distal and interstitial or proximal regions, whereas the last two had small chromosomes with early condensed chromatin in the proximal regions. Although they have been reduced in the literature to synonyms of B. pedatifida and B. villifolia , respectively, B. rubropunctata and B. purpureofolia should be treated as distinct species because of the differences in chromosome numbers and other chromosome features.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 513–517.     

Polymorphism of the Sphaerium corneum (Bivalvia, Veneroida, Sphaeriidae) revealed by cytogenetic and sequence comparison

Two populations of Sphaerium corneum were sampled from River Vilnelė and small pond in Vilnius, Lithuania. The chromosomes were studied using conventional Giemsa staining and karyometric analysis. Inter- and intra-individual variation in the diploid chromosome numbers was revealed and two different sources of chromosome variability were identified: B chromosomes and the structural changes of chromosomes of the basic (A) set. The chromosome set of the more common karyotypic form, 2 n  = 30, found in both populations, consists of all biarmed metacentric and meta-submetacentric chromosomes of gradually decreasing size. Small, biarmed, mitotically unstable B chromosomes were found in the cells of this karyotypic form. Specimens with 2 n  = 36 were found only in pond. No B chromosomes were detected in their cells. The karyotype is characterized by presence of two pairs of medium telocentrics and four pairs of small subtelocentrics. The remaining chromosomes are biarmed. Robertsonian fusions appear to be involved in formation of two karyotypic forms of S. corneum . DNA sequence analyses showed that ITS1 is identical in both karyotypic forms. On the other hand, differences in 16S sequence were revealed and two haplotypes, corresponding to two karyotypic forms, were identified. The present study opens new perspectives in establishing species-specific characters for confident identification of Sphaerium species and provides insights to the genetic intraspecific variability and possible mechanisms of speciation.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 53–64.     

Chromosome numbers in plant taxa endemic to the Balearic Islands

Mitotic chromosome numbers are reported from 20 vascular plant taxa that are endemic to the Balearic Islands and poorly known cytogenetically. The chromosome numbers of Arenaria grandiflora L. ssp. glabrescens (Willk.) G. López & Nieto Feliner (2 n  = 44), Dianthus rupicola Viv. ssp. bocchoriana L. Llorens & Gradaille (2 n  = 30), Solenopsis minuta (L.) C. Presl. ssp. balearica (E. Wimm.) Meikle (2 n  = 28), Romulea asumptionis Font Quer & Garcías Font (2 n  = 56), Scutellaria balearica Barceló (2 n  = 22) and Galium balearicum Briq. (2 n  = 22) have been determined for the first time. A new chromosome number was found in two populations of Helictotrichon crassifolium (Font Quer) Holub (2 n  =  c. 98) suggesting that this species is a high polyploid (14 x ), in contrast to an earlier report of a lower chromosome count (2 n  = 12 x  = 84). Cytogenetic observations suggest that Naufraga balearica Constance & Cannon has a diploid chromosome complement of 2 n  = 20, with 0–2 accessory chromosomes. The banded karyotype (chromomycin A₃) of Crepis triasii (Cambess.) Fries was determined for individuals belonging to eight populations from three islands. Two chromomycin A₃-positive regions were recorded in the diploid complement. These are present on the telomeric regions of the shortest subtelocentric chromosomes. This species is karyologically stable in chromosome number, karyotype and fluorochrome-banding pattern among populations from separate islands.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 219–228.     

Morphology of Pinus uliginosa (Pinaceae) needles from populations exposed to and isolated from the direct influence of Pinus sylvestris

Eleven needle traits of two contrasting populations of Pinus uliginosa ( P. mugo complex), one located in a Pinus sylvestris forest, the other isolated from the influence of this species, were compared to quantify the degree of hybridization. Statistically significant differences between the studied populations were found, but both of them differed by a similar degree from P. sylvestris . Surprisingly, the P. uliginosa population surrounded by the P. sylvestris forest, which was expected to consist of individuals with intermediate traits, appeared to be more closely related to P. mugo than those isolated from the direct influence of P. sylvestris . Large differences in the distances between the analysed pair of populations of P. uliginosa and P. sylvestris have not influenced the morphological traits of the species. Gene flow from P. sylvestris to P. uliginosa is possible, but is likely to be small and does not differ compared with populations of P. uliginosa . The decline observed during the last three decades in both populations of the species has not influenced their needle morphological traits.  © 2003 The Linnean Society of London , Botanical Journal of the Linnean Society , 2003, 142, 83–91.     

Karyological evolution in Rhaponticum Vaill. (Asteraceae, Cardueae) and related genera

Chromosome counts are reported in 29 populations representing 20 species and one subspecies of the genera Callicephalus (one sp.), Klasea (seven spp.), Myopordon (two spp.), Oligochaeta (one sp.), and Rhaponticum (nine spp., one ssp.). Eleven reports are new, the others supplement limited previous data. New and published cytological data are summarized and evaluated critically in light of current taxonomic treatments and evolutionary hypotheses. Basic chromosome numbers are a valuable source of taxonomic information and relate well to the phylogeny. They are quite conserved in Klasea (only x  = 15), the sister group of the remaining genera, and in the early branching genera of the Rhaponticum group ( x  = 14 for Centaurothamnus and Callicephalus ). However, a more complex pattern of genome evolution is present among the more derived clades, Oligochaeta plus Myopordon ( x  = 12, 14) and the genus Rhaponticum ( x  = 12, 13). The occurrence of x  = 12 in Oligochaeta divaricata and Rhaponticum carthamoides seems to be the result of a recent event of chromosome fusion, as suggested by the fragility of particularly long pairs of chromosomes. The use of chromomycin staining, including an enzymatic digestion step that facilitates spreading and permits a reduction of potential fragmentation, only partially resolves this problem.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 193–201.     

Chromosome variation in the genus Pinellia (Araceae) in China and Japan

The chromosome numbers of 23 populations belonging to seven species of Pinellia Tenore were counted. The basic chromosome number of all the species studied was x  = 13, and the previously reported basic numbers x  = 9 and x  = 14 in P. ternata and P. cordata were not confirmed. Chromosome numbers of 2 n  = 26 in P. polyphylla and 2 n  = 78 in P. integrifolia are reported for the first time. Heptaploid (7 x ) and nonuploid (9 x ) are new ploidy levels reported for P. ternata , and hexaploid for P. cordata. Taxonomic, phylogenetic and phytogeographical inferences are made for the genus. Particular attention was given to P. ternata , the most widely distributed species in the genus, and considerable variation of chromosome number was found in its different populations. Based on chromosome studies of 11 populations of P. ternata , together with 12 populations reported in previous studies, the lower reaches of the Yangze River are identified as its centre of origin. From there it dispersed, with generation of hexaploid, heptaploid, octoploid and nonuploid forms.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 449–455.     

Chromosome counts in Asian Artemisia L. (Asteraceae) species: from diploids to the first report of the highest polyploid in the genus

This work comprises 24 reports of chromosome numbers in 24 Artemisia L. species from Asia. Ten are included in the subgenus Dracunculus Besser and the rest belong to other subgenera. Seven counts are new reports, 14 are consistent with scarce previous ones, and three contribute new ploidy levels. That carried out in A. medioxima reports the highest ploidy level ever counted for the genus (16 x ). There is only one species with x  = 8 as the basic chromosome number. In the remaining x  = 9-based species, ploidy levels range from 2 x to 16 x , illustrating the great role played by polyploidy in the evolution of the genus.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 301–310.     

Ferulago trojana (Umbelliferae), a new species from western Turkey

Somatic chromosomes of four species of Ceratozamia , C. hildae , C. kuesteriana , C. mexicana and C. norstogii , and Stangeria eriopus , were observed and compared by the fluorescence in situ hybridization method using 5S ribosomal (rDNA) probes. The four Ceratozamia species and S. eriopus showed the same chromosome number of 2 n  = 16, and had similar karyotypes, comprising 12 metacentric (m), two submetacentric (sm) chromosomes and two telocentric (t) chromosomes. The four Ceratozamia species exhibited a proximal 5S rDNA site in the interstitial region of two m chromosomes. Stangeria eriopus exhibited a distal 5S rDNA site in the interstitial region of two m chromosomes, which probably indicates that the two genera differ in chromosome structure by at least one paracentric inversion. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 499–504.     

Karyological notes on eight species of Achillea L. (Asteraceae, Santolinoideae) from Turkey

The chromosome number and morphology of eight species belonging to the section Santolinoideae of the genus Achillea L. (Asteraceae) were investigated using karyological techniques. The species studied were Achillea cucullata (Hausskn.) Bornm., A. vermicularis Trin., A. monocephala Boiss. & Bal., A. boissieri (Hausskn.) Bornm., A. santolina L., A. gypsicola Hub.-Mor., A. goniocephala Boiss. & Bal., and A. spinulifolia Fenzl ex Boiss. In these species, the chromosome numbers were 2 n  = 18 in A. vermicularis , A. monocephala , and A. boissieri , 2 n  = 18A + 1B in A. spinulifolia , 2 n  = 36 in A. cucullata and A. santolina , and 2 n  = 54 in A. gypsicola and A. goniocephala . All of the chromosomes had median point (M), median region (m), or submedian (sm) centromeres. An increase in asymmetry was not observed in the karyotypes of the species.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 573–580.