Karyomorphology of the endangered Trigonobalanus doichangensis (A. Camus) Forman (Fagaceae) and its taxonomic and biogeographical implications

Cytological studies were carried out for the first time on five populations of Trigonobalanus doichangensis in China and Thailand. In all populations, the pattern of interphase nuclei was of the simple chromocentre type, the mitotic prophases were of the proximal interstitial type and chromosome numbers were 2 n  = 2 x  = 14. Two B chromosomes were commonly observed at prophase and prophase–metaphase, but rarely at metaphase. Karyotype variation among the populations at the diploid level was limited, but there were some distinguishing cytological characters. Based on the comparison of all the available data on cytology, taxonomy, phytogeography and molecular systematics related to the genus Trigonobalanus , we recommend that the three species of Trigonobalanus comprise the subfamily Trigonobalanoideae and that the genus should not be segregated into three monotypic genera.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 321–330.     

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.     

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.     

Chromosome numbers in Hieracium and Pilosella species (Asteraceae) from the Iberian Peninsula and the Balearic Islands

Mitotic chromosome numbers are reported from 16 Hieracium and two Pilosella species from the Iberian Peninsula and the Balearic Islands that are little known cytogenetically. The chromosome numbers of Hieracium aragonense Scheele (2 n  = 27, 36), H. compositum Lapeyr. (2 n  = 27), H. murcandidum G. Mateo (2 n  = 27), H. spathulatum Scheele (2 n  = 27), H. segurae Mateo (2 n  = 27), H. teruelanum Mateo (2 n  = 27), H. valentinum Pau (2 n  = 27), Pilosella pseudovahlii (De Retz) Mateo (2 n  = 18), and P. tardans (Peter) Sójak (2 n  = 36) were determined for the first time. New cytotypes were detected in H. cordifolium Lapeyr. (2 n  = 27) and H. loscosianum Scheele (2 n  = 36). The karyotype of the studied species was similar in overall morphology and comprised metacentric and submetacentric chromosomes; secondary constrictions (up to two) were detected in the chromosome complements of some species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 311–320.     

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.     

Cytotaxonomy of species of Vernonia, section Lepidaploa, group Axilliflorae (Asteraceae, Vernonieae)

Mitotic chromosomes of eight populations of six species of Vernonia sensu Baker (Asteraceae, Vernonieae, section Lepidaploa , group Axilliflorae ) were studied to assess the validity of maintaining the genus as proposed currently ( sensu Baker) or of splitting it into several smaller genera ( sensu Robinson). The species were collected in areas of 'cerrado' and 'campo rupestre' in southern and central-western Brazil. The chromosome numbers were 2 n  = 20–40 and karyotypic analysis showed a predominance of metacentric chromosomes, with some submetacentrics. B chromosomes were seen in one population of V. geminata . The chromosomes varied in length from 0.9 to 4.6 µm, with a total chromatin length of 29.7–45.0 µm and a symmetry index of 41.2–46.9%. The intrachromosomal asymmetry index varied from 0.13 to 0.29, and the interchromosomal asymmetry index varied from 0.15 to 0.21. The dispersion diagram showed that population 1 of V. geminata had the most asymmetrical karyotype. These results do not provide sufficient evidence for a conclusive cytotaxonomic discussion of Vernonia , mainly because of the impossibility of associating any infrageneric group with specific karyotypic characters, and also because of the small number of species analysed so far.  © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 99–108.     

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.     

Cytotaxonomy of Lychnophoriopsis Sch.Bip. and Paralychnophora MacLeish species (Asteraceae: Vernonieae: Lychnophorinae)

The genera Lychnophoriopsis and Paralychnophora are endemic to Brazilian 'campo rupestre' in the States of Minas Gerais and Bahia. To analyse the status of the cytotaxonomy of these genera, we carried out chromosome counts on several species. Two chromosome numbers were obtained for four species of Paralychnophora : 2 n  = 36 and 2 n  = 38. For three species of Lychnophoriopsis , only 2 n  = 36 was observed. The chromosome size (1.0–2.58 µm) and morphology (mainly metacentric, with some submetacentric) were analysed in four species of Paralychnophora and showed relative constancy .   © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 109–114.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Karyology of Limonium (Plumbaginaceae) species from the Balearic Islands and the western Iberian Peninsula

Somatic chromosome numbers, conventional karyotype features and idiograms are reported for 27 Limonium species inhabiting the Western Mediterranean basin (Iberian Peninsula and the Balearic Islands). The chromosome numbers of Limonium barceloi (2 n  = 36), L. ejulabilis (2 n  = 24), L. inexpectans (2 n  = 26), L. interjectum (2 n  = 24), and L. scopulorum (2 n  = 25) were determined for the first time. In addition, new aneuploid and/or polyploid cytotypes are reported in L. alcudianum (2 n  = 26), L. bonafei (2 n  = 26), L. camposanum (2 n  = 26) , L. companyonis (2 n  = 26), L. dufourii (2 n  = 26), L. geronense (2 n  = 36), L. marisolii (2n = 54), L. migjornense (2 n  = 50), and L. pseudodictyocladon (2 n  = 16). A group of polyploid species showed karyotypes comprising homologous chromosomes in groups of three ( L. antonii-llorensii, L. ejulabilis, L. interjectum, L. virgatum, and L. wiedmanii ) , four ( L. geronense ), or six ( L. marisolii ), which suggests an autopolyploid origin. Other polyploid species were characterized by the presence of two different chromosome sets ( x  = 8 and x  = 9) in the genome. The species L. alcudianum , L . bonafei , L. camposanum , L. companyonis , L. dufourii , L. gibertii , L. girardianum , L. inexpectans , L. leonardi-llorensii , L. magallufianum , L. migjornense , L. minoricense , and L. scopulorum showed various combinations of paired and unpaired x  = 8 and x  = 9 chromosome sets, suggesting that they are allopolyploids. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 257–272.     

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.     

Chromosomal evolution of South American frugivorous butterflies in the Satyroid clade (Nymphalidae: Charaxinae, Morphinae and Satyrinae)

We describe the chromosome numbers of a monophyletic group of Satyroid subfamilies of primary fruit-attracted butterflies from South America: Charaxinae, Morphinae (including Brassolini) and Satyrinae. The charaxines do not have a distinct modal number. Their chromosome numbers are in the range n  = 6–50, with n  = 7–9, n  = 12, n  = 16, n  = 19–21, n  = 26, and n  = 28–31 being the most common numbers. Within the Morphinae, the Morphini have a modal n  = 28 and the Brassolini a modal n  = 29, with few exceptions. The Neotropical satyrines, in particular the basal species, have a weak modal n  = 29, which is a strong modal number in Palearctic satyrines. The African satyrines have an equally strong modal n  = 28. Most Neotropical satyrines have, like charaxines, chromosome numbers lower than the weak modal n  = 29, and often half this modal, but there are genera with stable numbers among the satyrines and charaxines. Evidently, the Neotropical satyroids descend from basal Nymphalidae with the typical lepidopteran modal number of n  = 31, which have also given rise to the Heliconiini with modal n  = 31 and 21 and Ithomiinae with modal numbers of n  = 14–15.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 467–481.     

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.