Cytotaxonomical analysis of <Emphasis Type="Italic">Momordica</Emphasis> L. (Cucurbitaceae) species of Indian occurrence

Somatic chromosome number and detailed karyotype analysis were carried out in six Indian Momordica species viz. M. balsamina, M. charantia, M. cochinchinensis, M. dioica, M. sahyadrica and M. cymbalaria (syn. Luffa cymbalaria; a taxon of controversial taxonomic identity). The somatic chromosome number 2n = 22 was reconfirmed in monoecious species (M. balsamina and M. charantia). Out of four dioecious species, the chromosome number was reconfirmed in M. cochinchinensis (2n = 28), M. dioica (2n = 28) and M. subangulata subsp. renigera (2n = 56), while in M. sahyadrica (2n = 28) somatic chromosome number was reported for the first time. A new chromosome number of 2n = 18 was reported in M. cymbalaria against its previous reports of 2n = 16, 22. The karyotype analysis of all the species revealed significant numerical and structural variations of chromosomes. It was possible to distinguish chromosomes of M. cymbalaria from other Momordica species and also between monoecious and dioecious taxa of the genus. Morphology and crossability among the dioecious species was also studied. Evidence from morphology, crossability, pollen viability and chromosome synapsis suggests a segmental allopolyploid origin for M. subangulata subsp. renigera. The taxonomic status of the controversial taxon M. cymbalaria was also discussed using morphological, karyological and crossability data.     

Karyological studies on eight species of <Emphasis Type="Italic">Onobrychis</Emphasis> genus in Turkey

This study used karyological techniques to determine the chromosome numbers and morphology of eight species of Onobrychis L. (O. caput-galli (L.) Lam, O. aequidentata (Sibth. & Sm.) d’ Urv, O. fallax Freyn & Sint. var. fallax, O. lasiostachya Boiss, O. viciifolia Scop., O. oxyodonta Boiss. subsp. armena (Bois. & Huet) Aktoklu, O. hypargyrea Boiss. and O. cappadocica Boiss.). The results of this study determined the chromosome numbers of O. cappadocica as 2n = 16; O. viciifolia as 2n = 28 and the other species as 2n = 14 The karyotypes of species consisted of median-centromeric (m) or submedian-centromeric (sm) chromosomes. However, O. oxyodonta Boiss. subsp. armena (Bois. & Huet) Aktoklu was found to have only the median-centromeric (m) chromosomes. According to the results of the present study, of the eight Onobrychis taxa, only O. hypargyrea has a pair of satellite chromosomes (sat-chromosome). Furthermore, this study detected karyotype asymmetry.     

Karyotype variation within some native populations of oriental spruce (<Emphasis Type="Italic">Picea orientalis</Emphasis>) in Turkey

Karyological studies have been investigated within 8 native Anatolian populations of oriental spruce (Picea orientalis (L.) Link) in Turkey. The somatic chromosome number of 2n = 2x = 24 has been observed in all accessions. The karyotypes are generally asymmetrical with most of the chromosomes having median to median-submedian centromeres. Inter-population variability of the karyotype was summarized with cluster analysis. We found that the karyotypes have positively correlated with the altitudes of the natural habitats. The high value of karyotype asymmetry may be attributed to both microenvironment and natural regeneration methods used in oriental spruce.     

Karyoevelution on the genus Onosma L. (Boraginaceae): quantification of chromosome number heterogeneity

In this survey, chromosome counts of different species belonging to the genus Onosma are summarized and then karyological patterns available including frequency of cytotype occurrence, percentage of taxa with particular basic chromosome number and rate of polyploidy in the genus are evaluated. Quantitative parameters have been used to characterize chromosome number (CN) variation. In order to verify if variation patterns differ between three groups of Onosma, Index of CN Heterogeneity (ICNH) was quantified. In addition, meiotic chromosome numbers of 14 populations belonging to 11 species growing in Iran, namely Onosma araratica (2n = 2x = 16), O. asperrima (2n = 2x = 16), O. bulbotricha (2n = 2x = 18), O. kotschyi (2n = 2x = 16), O. microcarpa (2n = 2x = 16), O. nigricaulis (2n = 2x = 16), O. nervosa (2n = 2x = 16), O. obtusifolia (2n = 2x = 16), O. pachypoda (2n = 2x = 16), O. stenosiphon (2n = 2x = 20) and O. subsericea (2n = 2x = 16), were determined. With the exception of O. microcarpa and O. bulbotricha, all chromosome counts are reported for the first time. (© 2013 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Karyological analysis of the genus Canna (Cannaceae)

Somatic chromosome numbers and karyological features in 22 taxa of the genus Canna (Cannaceae) were investigated and are presented here for the first time. The chromosome numbers were primarily reported for C. bangii, C. indica var. sanctae-rosae and C. tulianensis. A triploid somatic chromosome number of 2n = 27 was found only in C. discolor, while the remaining taxa were diploids with 2n = 18. Chromosome numbers are considered to be remarkably stable within the Cannaceae. Canna shows a symmetric karyotype that consists mostly of median chromosomes, but with some subterminal or submedian chromosomes, and a karyotype homogenous in variation in chromosome length. The karyotype of some taxa of Canna, furthermore, is characterized by median chromosomes with a secondary constriction in the centromeric region. Cytotaxonomical characteristics of the family Cannaceae and the related groups in the Zingiberales are discussed.     

A simple parameter of dispersion index that serves as an adjunct to karyotype asymmetry

In order to refine the measure of karyotype asymmetry a new chromosomal parameter of dispersion index is proposed that has the potential to decipher even the minor karyotypic variations, thus permitting further evolutionary gradations to the karyotype asymmetry classes of Stebbins. The higher the dispersion index, the more specialized would be the karyotype. The dispersion index takes into account the variance lor gradual change in chromosome size within a complementvis-a-vis variance for the position of centromere in a karyotypic totality. The dispersion index is calculated as the proportionate measure of centromeric gradient to the coefficient of variation for chromosome length; wherin centromeric gradient = length of median short arm — length of median chromosome. Thus, the three most important karyotypic criteriaviz., differences in: absolute chromosome size, position of centromere and relative chromosome size, are all covered in the proposed parameter. The effectiveness of dispersion index has been tested on a plant taxa,Papaver L., where karyomorphological details, nuclear DNA content, and morphotaxonomic parameters have been amply elucidated from an evolutionary stand point. It is hoped that dispersion index would find immense utility in delimiting species interrelationships particularly in the closely related taxa, when applied in conjunction with other systematic parameters.     

Karyological studies of some representatives of <Emphasis Type="Italic">Tanacetum</Emphasis> L. (Anthemideae-Asteraceae) from north-east Anatolia

The karyotypes of nine Tanacetum taxa distributed in north-east Anatolia, Turkey, were determined and evaluated by cluster analysis and principal-components analysis. Chromosome numbers were 2n = 2x = 18 (8 taxa) and 4x = 36 (1 taxon). Somatic chromosome numbers of two taxa and a new ploidy level in one taxon are reported for the first time. Karyotype analysis indicated that chromosomes of Tanacetum taxa have predominantly median centromeres. The taxa studied differed significantly in the size of the short arms and long arms, and the arm ratio of each pair of homologous chromosomes, indicating structural rearrangements of the chromosomes have been involved in diversification of the taxa. They were placed in 2A, 3A, and 2B of Stebbins’ karyotype classification, showing the presence of a primitive symmetrical karyotype in the genus. Several systematic and evolutionary aspects of the genus are discussed on the basis of karyological data.     

Crocus ilvensis sp. nov. (sect. Crocus,Iridaceae), endemic to Elba Island (Tuscan Archipelago,Italy)

A new species of Crocus sect. Crocus is described as an endemic of Elba Island (Tuscan Archipelago, Italy): C. ilvensis Peruzzi & Carta sp. nov. (2n=8). The new species was wrongly referred to C. corsicus Vanucchi (2n=18), C. etruscus Parl. (2n=8) or C. vernus Hill. subsp. vernus (2n=8, 16) by previous authors. Its karyotype structure, asymmetry and chromosome dimensions show affinity with C. etruscus, and no clear relations with other related taxa. From a morphological point of view, C. ilvensis appears intermediate between C. etruscus and C. vernus subsp. vernus, showing however a peculiar combination of character‐states. The new species is completely allopatric with other Crocus sect. Crocus taxa and its possible role in the origin of tetraploid races of C. vernus is discussed.     

Towards resolving the systematics of Cerastium subsection Cerastium (Caryophyllaceae): a cytogenetic approach

Analyses of mitotic chromosome numbers and nuclear DNA content were performed for 39 populations of 17 perennial Cerastium taxa from south‐eastern Europe. The DNA content ranged from 2C = 2.43 to 8.78 pg, revealing four ploidy levels corresponding to 4x (2n = 36), 8x, 12x and 16x. High‐polyploid cytotypes with a greater range of ploidy (up to 2n = 144) occur mostly in the central mountainous parts of the Balkan Peninsula. The chromosome number was determined for the first time for C. dinaricum (2n = 36 + 1B), C. decalvans subsp. orbelicum (2n = 36), C. decalvans subsp. glutinosum (2n = 36), C. neoscardicum (2n = 144), C. malyi subsp. serpentini (2n = 144) and C. moesiacum s.s. (2n = 144). New chromosome counts were recorded for C. arvense subsp. strictum (2n = 108), C. banaticum subsp. kosaninii (2n = 36) and C. grandiflorum (2n = 36). For the first time, flow cytometry was used to estimate C values for six species (15 taxonomic entities). The intraspecific variation quotient of C values is high, ranging from 1.003 in C. malyi to 1.306 in C. decalvans subsp. decalvans. The variation in chromosome size among both tetra‐ and octoploid members of Cerastium is much more prominent than in most other angiosperm polyploid series. Significant genome downsizing after polyploidization was observed in some investigated taxa. Differences in ploidy levels and monoploid genome size values confirm the taxonomic status of C. decalvans subsp. glutinosum and C. decalvans subsp. leontopodium. The results obtained indicate a possible close relationship between C. banaticum and C. grandiflorum, but not C. arvense. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 182 , 205–224.     

Karyological studies in IberianSonchus (Asteraceae: Lactuceae): S. oleraceus, S. microcephalus andS. asper and a general discussion

Chromosome numbers and karyotypes ofSonchus oleraceus (n=16, 2n=32),S. asper (n=9, 2n=18 for subsp.asper and subsp.glaucescens, andS. microcephalus (n=15, 2n=30) are studied; for the first time their idiograms are given. Karyotypes mainly comprise small chromosomes with a degree of asymmetry 2B inS. oleraceus andS. microcephalus and 2A or 2A-2B inS. asper. No karyological differences can be pointed out betweenS. asper subspecies. Data presented support the amphiploid character ofS. oleraceus, and the origin ofS. microcephalus through a dysploid process involving the former taxon. In Iberian representatives of the genus, diagrams of karyotype asymmetry indices show a cluster grouping for species, with the detached exceptions ofS. maritimus andS. crassifolius, which spontaneously hybridize in central Spain. A review of available karyological data shows that in the evolution of the genusSonchus s.l. and relatives, the basic chromosome number x=9 has generally been maintained. Dysploidy is restricted to the seriesS. bourgeaui (n=8) —S. tenerrimus (n=7) andS. oleraceus (n=16) —S. microcephalus (n=15), evolutionarily related and included in the present subgenusSonchus. Polyploidy has been detected in a total of nine taxa ofSonchus and in the generaEmbergeria, Kirkianella, andDendroseris, being more common in peripheral regions of the distribution area of the group. Five diversification centers are proposed for the whole group, of which the Western Mediterranean area, including the Iberian Peninsula, is related to diversification of the present subgenusSonchus.     

Some evidence for infrageneric classification in Ziziphora L. (Lamiaceae:Mentheae)

The genus Ziziphora L. (Lamiaceae) consists of annual herbs, except Ziziphora clinopodioides, and is represented by six taxa in the flora of Turkey: Z. clinopodioides, Z. capitata, Z. persica, Z. tenuior, Z. taurica subsp. taurica and Z. taurica subsp. cleonioides. They are strong aromatic herbs and important medicinal plants. This study has been conducted to determine the palynological and karyological features of these taxa. The pollen morphology and exine structure of Ziziphora species were investigated by light microscope and scanning electron microscope. Pollen grains were determined as hexacolpate, medium in size and a circular or elliptic amb. The shape of pollen grains varies from oblate to prolate spheroidal. Ornamentation types of pollen grains are bireticulate, microreticulate or perforate. Somatic chromosome number has been determined and counted as 2n = 16, 18. Results indicate that the pollen characters and chromosome numbers of the genus are valuable for taxonomic applications and may be useful for infrageneric classification.     

Cytotaxonomy of Prenanthes faberi (Compositae–Cichorieae)

The generic placement of Prenanthes faberi Hemsl. (Compositae–Cichorieae) has long been a controversial matter. Recent molecular phylogenetic analyses indicate its inclusion in the genus Faberia Hemsl. The present cytological investigation shows that this species has a chromosome number of 2n = 34 and a fairly symmetric karyotype consisting of 26 median centromeric (m) and 8 submedian centromeric (sm) chromosomes varying in length between 1.72–4.20 μm. The karyological characters of P. faberi are remarkably consistent with those reported previously in species of Faberia. The transference of this species to Faberia is strongly supported.     

Karyosystematic studies on threeCrepis species (Asteraceae) endemic to Greece

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

Comparison of the Giemsa C-banded karyotypes of the three subspecies ofPsathyrostachys fragilis,subspp.villosus (2x),secaliformis (2x, 4x), andfragilis (2x) (Poaceae), with notes on chromosome pairing

The karyotypes of diploidP. fragilis subsp.villosus (2n = 2x = 14) and tetraploid subsp.secaliformis (2n = 4x = 28) were studied by Giemsa C- and N-banding, and AgNO₃ staining and compared with the karyotype of subsp.fragilis (2x). The complements of subsp.villosus and subsp.fragilis were similar, with 8 metacentric and 6 SAT-chromosomes, one metacentric and two submetacentric pairs, with small to minute, polymorphic, heterochromatic satellites. The complement of subsp.secaliformis on the whole agreed with a doubling of the complement of diploidP. fragilis, suggesting autopolyploidy. Only the presence of 12 nucleoli in interphases identified 6 SAT-chromosome pairs. In subsp.villosus one or two extra micronucleoli indicated a chromosome pair with very low nucleolusforming activity, bringing the number of SAT-chromosome pairs to 4. This number may be a characteristc ofPsathyrostachys. Besides very small, inconsistently observed bands, the C-banding pattern consisted of 0–3 small bands per chromosome at intercalary and terminal locations, and at NORs. The level of banding pattern polymorphism was low, but enough to indicate that the taxa are outbreeders. Similarities in chromosome morphology and C-banding patterns identified homology of all chromosomes of subsp.villosus, but for 12 pairs only in subsp.secaliformis. Between plants, reliable identification of homology and homoeology (subsp.secaliformis) was possible only for the SAT-chromosomes and the shortest metacentrics. Chromocentres were very small and the amount of constitutive heterochromatin was low. N-banding stained chromosomes uniformly. The basic karyotypes of theP. fragilis taxa were similar to those ofP. juncea, P. lanuginosa, andP. stoloniformis supporting a close relationship and the presence of a common genome, N. NORs had different nucleolus-forming activities. Meiotic analysis demonstrated a high level of bivalent pairing in the three taxa. A chromosomal rearrangement was suggested in subsp.villosus. The low multivalent frequency in subsp.secaliformis indicates the presence of a pairing regulation mechanism. The majority of chiasmata were interstitial. Pollen grain size discriminated between diploid and tetraploid taxa. The existence of a diploid cytotype of subsp.secaliformis is supported by pollen measurements of herbarium material.     

HETEROCHROMATIN BANDING IN BOYKINIA,HEUCHERA, MITELLA,SULLIVANTIA, TIARELLA,AND TOLMIEA (SAXIFRAGACEAE)

Karyotypic differences were sought among species of Boykinia, Heuchera, Mitella, Sullivantia, Tiarella, and Tolmiea utilizing a modification of the Hy-banding technique. Prominent centromeric and some telomeric heterochromatin banding was observed. Boykinia aconitifolia and species of Sullivantia possess an identical banded karyotype, while four species of Heuchera, Mitella diphylla, Tiarella cordifolia, and Tolmiea menziesii (the latter at the tetraploid level) are characterized by a second, slightly different banded karyotype. In Sullivantia, Giemsa C-banding stains the same chromosomal regions revealed by Hy-banding. Larger amounts of heterochromatin are present in chromosomes of species of Heuchera, Mitella, Tiarella, and Tolmiea than in chromosomes of Sullivantia species and Boykinia aconitifolia. These karyological observations confirm generic relationships and demonstrate the systematic applicability of chromosome banding techniques to plants with very small chromosomes.     

Chromosome painting in<Emphasis Type="Italic"> Callicebus lugens</Emphasis>, the species with the lowest diploid number (2<Emphasis Type="Italic">n</Emphasis>=16) known in primates

Cytogenetic studies have shown that New World primates are karyologically diverse and highly derived. The genus Callicebus is the best example of this karyological diversity, with diploid numbers ranging from 2n=50 to 2n=16. We report on Callicebus lugens, which has the lowest diploid number (2n=16) yet found in the primate order and represents a striking example of extreme karyotypic shuffling. To better understand the genomic rearrangements that have resulted in this extremely low diploid number, we mapped chromosome homologies between C. lugens and humans by in situ hybridization. The total number of hybridization signals was 42, excluding the Y chromosome, with a total of 34 syntenic associations not found in humans. This species has one of the most derived karyotypes among the Platyrrhini. Fusion has been the predominant mode of karyological evolution, although fissions and inversions have also transformed the C. lugens karyotype. Remarkably in such a highly rearranged karyotype, the synteny of 11 human chromosomes (4, 5, 9, 12, 13, 14, 17, 18, 20, 21, and X) was maintained intact, even if most of these human-homologous gene clusters were translocated. Other human syntenies, such as homologues to human chromosomes 10 and 16, were highly fragmented. Comparisons of the C. lugens-human homology map with those of other New World primates have not yet helped establish a phylogenic arrangement between congeneric species or link Callicebus with any other genus.Communicated by S. Henikoff     

A cytological study on Kelloggia (Rubiaceae), an intercontinental disjunct genus between eastern Asia and western North America

A cytological study was carried out for the first time on four populations of the only two species of Kelloggia (Rubiaceae), which occur disjunctly in eastern Asia (K. chinensis Franch.) and western North America (K. galioides Torr.). The consistent mitotic prophase chromosome condensation pattern and interphase nuclei type were determined for both species. The chromosome base number of the genus is suggested to be x=11. The karyotype of 2n=22=2x=16m+6sm was examined for both species. The karyotypical asymmetry of 1A and 2A was found in K. chinensis and K. galioides, respectively. According to the predominant evolutionary direction of karyotype asymmetry in angiosperms, K. galioides (2A) seems slightly more evolved than K. chinensis (1A). Our finding is consistent with the hypothesis of the Old World origin of Kelloggia based on molecular study.     

The phylogenetic position of Apterosperma (Theaceae) based on morphological and karyotype characters

Classifications of Theaceae have usually placed the endangered monotypic genus Apterosperma in tribe Schimeae (x=18), whereas recent molecular phylogenetic evidence supports its transfer to tribe Theeae (x=15). Molecular data have not resolved the phylogenetic position of Apterosperma within Theeae. We investigated the chromosome number and karyotype of Apterosperma in the context of molecular and morphological phylogenetic evidence to provide further insight into the placement of Apterosperma within Theaceae. The chromosome number and karyotype was found to be 2n = 30 = 26m + 4sm, consistent with the transfer of Apterosperma to tribe Theeae. When the chromosome data were incorporated into a data set of 46 other nonmolecular characters, Apterosperma was placed as the first-diverging lineage within the clade comprising tribe Theeae. This supports its placement based on molecular data. The low intrachromosomal asymmetry (type 1A) of Apterosperma, presumably ancestral for the family, is also consistent with this placement. Character optimization strongly supports a base chromosome number of x=15 for tribe Theeae. Because of variable and sometimes conflicting chromosome count reports of species in tribes Schimeae and Stewartieae, the base chromosome number of Theaceae could be either x=15 or 17.     

Chromosome evolution in pseudoxyrhophiine snakes from Madagascar: a wide range of karyotypic variability

In this first cytogenetic survey on the lamprophiid snake subfamily Pseudoxyrhophiinae, we studied the karyology of ten snake species belonging to seven genera from Madagascar (Compsophis, Leioheterodon, Liophidium, Lycodryas, Madagascarophis, Phisalixella and Thamnosophis) using standard and banding methods. Our results show a wide range of different karyotypes ranging from 2n = 34 to 2n = 46 elements (FN from 40 to 48), with nucleolus organizer regions (NORs) on one (plesiomorphic) or two (derived/apomorphic) microchromosome pairs, and W chromosome at early or advanced states of diversification from the Z chromosome. The observed W chromosome variations further support the most accepted hypothesis that W differentiation from the Z chromosome occurred by progressive steps. We also propose an evolutionary scenario for the observed high karyotype diversity in this group of snakes, suggesting that it is derived from a putative primitive pseudoxyrhophiine karyotype with 2n = 46, similar to that of Leioheterodon geayi, via a series of centric fusions and inversions among macrochromosomes and translocations of micro‐ either to micro‐ or to macrochromosomes. This primitive Pseudoxyrhophiinae karyotype might have derived from a putative Lamprophiidae ancestor with 2n = 48, by means of a translocation of a micro‐ to a macrochromosome. In turn, the karyotype of this lamprophiid common ancestor may have derived from the assumed primitive snake karyotype (2n = 36 chromosomes, with 16 biarmed macro‐ and 20 microchromosomes) by a series of centric fissions and one inversion. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 450–460.