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.     

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.     

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.     

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 studies in Tripleurospermum (Asteraceae, Anthemideae) from north-east Anatolia

Chromosome number and morphology in 14 taxa belonging to 19 populations of Tripleurospermum Sch. Bip. were studied using karyological and numerical taxonomical techniques. Data on chromosome measurements were analysed using cluster analysis. Chromosome numbers of these taxa are 2 n  = 2 x  = 18, 4 x  = 36 and 5 x  = 42–48. Seven records are new, two are not consistent with previous counts, and the remainder confirm the very limited previous data (one to three records). A new ploidy level (pentaploidy) is reported for the first time for the genus. Some correlations between ploidy levels and morphological characters are noted and several systematic and evolutionary aspects of the genus are discussed in the light of karyological data.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 146 , 427–438.     

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.     

Weintrauboa, a new genus of pimoid spiders from Japan and adjacent islands, with comments on the monophyly and diagnosis of the family Pimoidae and the genus Pimoa (Araneoidea, Araneae)

The spider genus Weintrauboa new genus (Araneae, Pimoidae) is described to place two species of pimoids from Japan and adjacent islands that were formerly classified in the linyphiid genus Labulla . Weintrauboa contortipes (Karsch) new comb., the type species, and W. chikunii (Oi) new comb. are redescribed. Parsimony analysis of morphological characters provides robust support for the monophyly of the genus Weintrauboa and corroborates the monophyly of Pimoa , Pimoidae, and the clade Linyphiidae plus Pimoidae. New diagnoses for Pimoa and Pimoidae are provided.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society , 2003, 139 , 261–281     

New chromosome counts in the Centaurea Jacea group (Asteraceae, Cardueae) and some related taxa

Twenty-seven chromosome counts are reported in 23 species of the genus Centaurea , mostly eastern endemic species of the Jacea group, which has become the core of the re-defined genus Centaurea . Twenty reports are new, one is a correction of a previous count, one is a confirmation of limited previous data and one represents a new basic number in the Centaureinae. The prevalence of the basic chromosome number x  = 9 among the Eastern sections of the Jacea group is confirmed, together with the close correlation between karyological data and classification of the genus. Two alternative hypotheses on the aberrant chromosome number (for the Centaureinae) found in C. behen are proposed. One of them, if verified, would confirm that a cycle of polyploidy and descending dysploidy is a key mechanism in the speciation of the group.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 345–352.     

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.     

Chromosomal studies in species of Salvia (Lamiaceae) from Argentina

Meiotic and mitotic chromosomes of 13 species of Salvia from Argentina were studied. Most of our data are new counts but some, mainly from cultivated species, confirm previous reports or represent numbers which are different from those cited previously. Only Salvia gilliesii Benth. and the introduced species S. coccinea Juss., S. farinacea Benth., S. involucrata Cav. and S. microphylla Kunth are diploids. S. cardiophylla Benth., S. procurrens Benth. , S. splendens Roem. & Schult. and S. uliginosa Benth. are tetraploids; S. stachydifolia Benth. is hexaploid; and S. guaranitica A. St.-Hil., S. pallida Benth. and S. rypara Briq. are octoploids. The basic number most frequently found is x  = 11, but two species, S. procurrens and S. uliginosa , have x  = 13. S. farinacea has x  = 10 but this species is a widespread cultivated plant, not native in this area. Seven of the studied species showed one to three B chromosomes. We discuss some systematic and evolutionary aspects of the genus in the light of the cytogenetic data, the relationships between geographical distribution and chromosome numbers in relation to levels of ploidy and basic chromosome numbers, and make comparisons with some of the different taxonomic sections.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 483–490.     

Historical biogeography of Hexabathynella , a cosmopolitan genus of groundwater Syncarida (Crustacea, Bathynellacea, Parabathynellidae)

Hexabathynella is the only cosmopolitan genus of the order Bathynellacea (Crustacea). The known species number 18, found in Europe (9), Africa (1), South America (2), North America (3) and Australia and New Zealand (3). Phylogenetic analysis suggests that the least derived species are those from South America and the most derived those from the Iberian Peninsula, North America and Australia. The five species with the most plesiomorphic characters occur in salt or brackish water, which supports a marine origin for the genus. Phylogenetic and biogeographical analyses suggest that the distribution of the genus can be explained by dispersion and a double vicariant biogeographical model based on plate tectonics and the evolution of the Tethys during the Mesozoic and Cenozoic.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 457–466.     

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.     

Evolution and phylogeny of the genus Natrix (Serpentes: Colubridae)

Some aspects of the natural history of snakes of the colubrid genus Natrix have been well studied. With their extensive European distribution and relative abundance, their ecology, reproduction and behaviour are well known. Yet other facets of their biology remain poorly understood. These include knowledge of Natrix phylogeny, hypotheses explaining the current distribution of the three extant members of the genus, and their evolution and relationships. In this study we used molecular data, the nucleotide sequences of four protein-coding mitochondrial genes (3806 bp total), to provide a well-supported phylogeny for the genus Natrix . With these molecular data, evidence from the fossil record, and knowledge of palaeogeological events, we used two approaches in designing a time scale which we used to date the major events in Natrix speciation and intraspecific variation. Our data strongly support a phylogeny for the genus in which N. maura is basal with N. natrix and N. tessellata being sister species. The calibrated molecular clock suggests that N. maura diverged from the common ancestor of the three species 18–27 mya and that N. natrix and N. tessellata diverged 13–22 mya. Although the ranges of these estimates are large they support an early Miocene to late Oligocene origin for the three species. Intraspecific divergence is estimated to have commenced 5.3, 6.0 and 6.7 mya with evolutionary rates of 1 : 1.25 : 1.35% per million years for N. maura, N. natrix and N. tessellata , respectively.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 127–143.     

Uniformity of karyotypes in Ligularia (Asteraceae: Senecioneae), a highly diversified genus of the eastern Qinghai–Tibet Plateau highlands and adjacent areas

Ligularia , a highly diversified genus in the eastern Qinghai–Tibet Plateau and adjacent areas, was chosen as a suitable subject in which to study speciation patterns in this 'hot spot' area at the chromosomal level. Chromosome numbers and karyotypes were studied in 23 populations of 14 species, most of which are endemic to this area. The basic number x  = 29 was confirmed for all species. Ligularia virgaurea was found to have diploid and triploid cytotypes, 2 n  = 58 and 87. Other species are only diploid, with 2 n  = 58. The karyotypes of all populations within any species, and all species spanning most sections and covering most of the morphological range in Ligularia , are very similar to each other, belonging to type 2A according to Stebbin's classification. This karyotype was also found in its close allies, e.g. Cremanthodium , Ligulariopsis , Parasenecio , and Sinacalia . Aneuploid reduction of chromosome number from 2 n  = 60 to 58 and karyotypic variation was found in Ligularia and its allies. Such a chromosomal pattern with few polyploids infers that variation of karyotype structure at the diploid level seems to be the predominant feature of chromosomal evolution in this group and sympatric speciation via hybridization and polyploidization has played a minor role in its species diversity.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 329–342.     

Seed micromorphology in the genus Neotinea Rchb.f. (Orchidaceae, Orchidinae)

The ultrastructure of testa seed in the genus Neotinea (Orchidaceae, Orchidinae) was examined for the first time. The morphology of the seed and of the anticlinal and periclinal walls was analysed by scanning electron microscopy. Quantitative data concerning the length and width of the seed and embryo, seed and embryo volume, free air space, and number of cells along the longitudinal axis are presented. In all species, the seeds are fusiform in shape with transverse ridges on the inner periclinal walls. This ornamentation pattern is characteristic for the genus Neotinea . It is a good diagnostic value supporting the monophyly of this genus, which has recently been proposed by several authors.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 133–140.     

Chromosome numbers in the tribes Anthemideae and Inuleae (Asteraceae)

Twenty-two chromosome counts of 19 taxa (21 populations) in the tribe Anthemideae and one member (one population) of the tribe Inuleae of the family Asteraceae are reported. The Anthemideae studied belong to the subtribes Artemisiinae (14 Artemisia taxa, and one species each of the genera Dendranthema , Filifolium and Neopallasia ) and Tanacetinae (one species each of the genera Lepidolopha and Tanacetopsis ). From the Inuleae, we studied one Inula species. Five counts are new reports (including two at generic level), six are not consistent with previous counts and the remainder are confirmations of very limited (one to four records) previous data. Most of populations of Anthemideae studied have the basic chromosome number x  = 9, with ploidy levels ranging from 2 x to 10 x . Dysploidy is also present, with two x  = 8 diploid taxa. The species of Inuleae studied is a diploid with x  = 10, also indicating dysploidy, other members of the same genus Inula having basic numbers of x  = 9 or 8.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 77–85.     

Chromosome study of the fern genus Cyrtomium (Dryopteridaceae)

Cyrtomium (Dryopteridaceae) is an Asiatic genus comprising about 40 species characterized by having anastomosing veins that form areolae with included veinlets. Twelve chromosome counts representing 11 taxa were made in the present study. The chromosome numbers of six species endemic to China are reported for the first time: C. uniseriale , 2 n  = 164; C. guizhouense , 2 n  = 82; C. shingianum , n  = 82, 2 n  =  c .164; C. chingianum , 2 n  = 164; C. urophyllum , 2 n  = 82 and C. aequibasis , 2 n  = 123. The results were used to show that (1) C. uniseriale should be reduced to varietal status under C. balansae ; (2) subseries Balansana is not closely related to any other Cyrtomium species and should be separated from Cyrtomium ; and (3) C. guizhouense and C. lonchitoides are basal groups when subseries Balansana is not considered. These results were also supported by morphology. A high level of differentiation is a distinct characteristic in Cyrtomium . The genus has two reproductive types: sexual and apogamous. Sexual species are distributed mostly in China. Southwestern China is a modern diversity centre of Cyrtomium , with 80.6% of known species occurring there. Many species are endemic, most of which are sexual tetraploids or apogamous triploids. Both the apogamous reproductive type and polyploid species are adaptions to limestone habitats.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 221–228.