Cytological studies on the Sino-Himalayan endemic Anisodus and four related genera from the tribe Hyoscyameae (Solanaceae) and their systematic and evolutionary implications

Cytological studies were carried out on eight species of five genera ( Anisodus , Atropanthe , Hyoscyamus , Mandragora and Przewalskia ) in the tribe Hyoscyameae (Solanaceae). First chromosome counts were reported in six species: Anisodus luridus , A. acutangulus , A. tanguticus , A. carniolicoides , Atropanthe sinensis and Mandragora caulescens , all with 2 n  = 8 x  = 48. Two records, for Hyoscyamus niger (2 n  = 2 x  = 34) and Przewalskia tangutica (2 n  = 4 x  = 44), were also confirmed. All species studied showed the proximal type of mitotic prophase chromosome condensation pattern. Three types of interphase nuclei were recognized: the round prochromosome type for Anisodus , Atropanthe and Mandragora , the rod prochromosome type for Przewalskia and the complex chromocentre type for Hyoscyamus . The cytological data supported the close relationship of all four species of Anisodus . Evidence from the interphase nuclei and chromosome base numbers supported the traditional classification of Hyoscyameae into two groups, i.e. Physochlaina praealta + Hyoscyamus (complex interphase type and x  = 7, 14, 17) and Przewalskia + Atropanthe + Anisodus + Scopolia + Atropa + Mandragora (prochromosome type and x  = 6, 11). Polyploidy is found in most species of the tribe in the Himalayan–Hengduan Mountains, as well as in the Mediterranean region, west–central Asia and eastern Asia. It seems that it probably occurred very early in Hyoscyameae evolution, before the uplift of the Himalayan–Hengduan Mountains. The Himalayan orogeny might have played a minor role in the polyploid evolution of plants in this tribe.  © 2005 The Linnean Society of London. Botanical Journal of the Linnean Society , 2005, 147 , 457–468.     

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.     

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.     

New or rarely reported chromosome numbers in taxa of subtribe Artemisiinae (Anthemideae, Asteraceae) from Mongolia

This study encompasses 25 chromosome counts of 18 species in the subtribe Artemisiinae (tribe Anthemideae) of the family Asteraceae, from Mongolia. Most (15 species) belong to Artemisia , the largest genus of the subtribe, whereas the others come from two genera very closely related to it: Ajania (two species) and Neopallasia (one species). Eleven counts are new reports, three are not consistent with previous reports and the remainder confirm scanty earlier information. The majority of the species have x  = 9 as their basic chromosome number, but there are some taxa with x  = 8. Ploidy levels range from 2 x to 6 x . The presence of B-chromosomes was detected in Ajania fruticulosa .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 203–210.     

The gynostemium of Hemipiliopsis purpureopunctata and Senghasiella glaucifolia, two taxonomically disputed species of Habenariinae (Orchidaceae)

The gynostemium structure and ontogeny of two taxonomically disputed orchids, Hemipiliopsis (= Habenaria ) purpureopunctata and Senghasiella (= Habenaria ) glaucifolia , are described and illustrated by scanning electron micrographs. The early gynostemium ontogeny of Hemipiliopsis purpureopunctata is shown to be fundamentally similar to that of the species of the tribe Orchideae that have been previously studied. This includes the initiation sequence of sepals, petals and lip, form and orientation of anthers, three-lobed condition of median carpel apex, and presence of auricles and basal bulges. During the later developmental stages some differences occur. The stigma processes of Senghasiella glaucifolia are united into a tongue-shaped organ, and the lateral rostellum lobes of Hemipiliopsis purpureopunctata protrude forwards with their viscidia positioned above the spur-mouth. Based on gynostemium characters, the generic rank of Hemipiliopsis was confirmed, but that of Senghasiella was not supported.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 191–196.     

Karyomorphology of Incarvillea (Bignoniaceae) and its implications in distribution and taxonomy

The karyomorphology of 11 species of the genus Incarvillea Juss. is reported. The chromosome numbers of all species studied are 2 n  = 22. The interphase nuclei and prophase chromosomes were found to be of the simple chromocentre type and the interstitial type, respectively. The asymmetry of the karyotype of I. arguta (two populations) in subgenus Amphicome is type 2A. The karyotypes of Incarvillea s inensis var. sinensis , I.  s inensis var. przewalskii, and I. olgae in subgenus Incarvillea are of asymmetry type 3A. The remaining nine species and one variety in subgenus Pteroscleris are also of asymmetry 3A. Data on three species and one variety studied are first reports. This study indicates that karyotype variation at the diploid level appears to be the predominant feature of chromosome evolution in the genus Incarvillea . According to this study of karyomorphology, morphological characteristics and geographical distribution, it seems that the three subgenera should be regarded as three independent genera. The geography of the genus is discussed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 113–121.     

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 numbers in the tribe Anthemideae (Asteraceae) from north-east Anatolia

Twenty-two chromosome counts of 19 taxa in the tribe Anthemideae of the family Asteraceae are reported from north-east Anatolia, Turkey. The taxa belong to the subtribes Achilleinae (four Achillea taxa), Anthemidinae (five Anthemis taxa), Artemisiinae (two Artemisia species), Leucantheminae (one Lecanthemum species), Matricariinae (two Tripleurospermum species) and Tanacetinae (five Tanacetum taxa). Six counts are new reports, seven are not consistent with previous counts, and the remainder are confirmations of very limited previous data. Most of the populations of Anthemideae studied have the basic number x  = 9, with ploidy levels ranging from 2 x to 7 to 8 x , but dysploidy is also present, with one diploid species, Artemisia austriaca , having x  = 8.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 203–211.     

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.     

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.     

Chromosomes of four species in three genera of Commelinaceae from China and their systematic implications

Chromosomes of four species in three genera of Commelinaceae from China were examined. Spatholirion longifolium had 2 n  = 20. This small genus is considered to be of a possible polyploid origin based on x  = 5, a basic number shared by its closest ally, the monospecific genus Streptolirion , with its only species having 2 n  = 10. The two genera were found to be very different from each other in their karyotypic constitution. The differences in both chromosome number (2 n  = 20 vs. 10) and karyotypic constitution support their separation as two independent genera. The remaining three species, Amischotolype hispida , Porandra ramosa and P. scandens , all had 2 n  = 36. In view of the occasional occurrence of 2 n  = 18 in the African Coleotrype , a genus very close to Amischotolype and Porandra in gross morphology, these three species are all very probably polyploid based on x  = 9. The high degree of karyotypic similarity of the three genera also strongly indicates their close affinity and supports their placement in the subtribe Coleotrypinae. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 399–403.     

Contribution to the karyological knowledge of Echinops (Asteraceae, Cardueae) and related genera

Twenty-six chromosome counts were made of several genera of the tribe Cardueae from various European and Asian provenances: Acantholepis (one species studied), Amphoricarpus (1), Chardinia (1), Echinops (14 species, 15 populations), Siebera (1), Staehelina (3) and Xeranthemum (4). Eleven of the reports are made for the first time, ten confirm previous counts, while the remainder report disparities with earlier records. The existence of different basic chromosome numbers and ploidy levels suggests dysploidy and polyploidy as the main mechanisms of chromosomal evolution in the taxa considered.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 337−344.     

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.     

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.     

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

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

Cytogenetics and chromosome banding patterns in Caesalpinioideae and Papilionioideae species of Pará, Amazonas, Brazil

Despite the abundance of woody legumes in Brazilian Amazonian rain forests, there are few chromosome counts on the native species of this important region. The present work presents such data for 13 species of Caesalpinioideae (the genera Bauhinia , Caesalpinia , Cassia , Chamaecrista and Senna ) and Papilionioideae ( Bowdichia , Centrosema and Dioclea ) collected from 17 natural populations. Our report represents the first chromosome counts for the genera Bowdichia and Dioclea and for four of the studied species. Observations are made on chromosome morphology, size, condensing behaviour and interphase nucleus structure. Banding with fluorochromes carried out for the first time in Caesalpinioideae revealed discrete CMA⁺/DAPI^– terminal (GC-rich) bands on 2–4 chromosome pairs of most species analysed, with a few species presenting discrete CMA^–/DAPI⁺ (AT-rich) bands. Significant differences in chromosome size, morphology and condensing behaviour were observed among members of the controversial tribe Cassieae ( Cassia , Chamaecrista and Senna ), revealing the tribe to be a heterogeneous group from the karyological point of view.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 181–191.     

Chromosome studies on some Asteraceae from South America

Mitotic or meiotic chromosome numbers for 42 accessions belonging to 39 species of different genera of Asteraceae were determined. First chromosome counts are reported for one genus ( Gymnocoronis ), 14 species, and one variety. These are as follows: Solidago chilensis var. megapotamica (2 n  = 2 x  = 18), Chromolaena barbacensis (2 n  = 3 x  = 30), Chromolaena christieana (2 n  = 3 x  = 30), Chromolaena hirsuta (2 n  = 4 x  = 40), Chromolaena verbenacea ( n  = 20 II, 2 n  = 4 x  = 40), Disynaphia multicrenulata (2 n  = 2 x  = 20), Gymnocoronis spilanthoides var. subcordata (2 n  = 2 x  = 20), Mikania thapsoides (2 n  = 4 x  = 68), Stevia commixta (2 n  = 2 x  = 22), Porophyllum brevifolium (2 n  = 4 x  = 44), Viguiera rojasii (2 n  = 2 x  = 34), Pterocaulon angustifolium (2 n  = 2 x  = 20), Gochnatia haumaniana (2 n  = 4 x  = 44), Senecio ostenii (2 n  = 4 x  = 40), Senecio pinnatus (2 n  = 8 x  = 80), and Lepidaploa amambaia (2 n  = 2 x  = 28). Chromosome numbers differing from those reported previously in the literature were found in Campuloclinium macrocephalum (2 n  = 2 x  = 20), Melanthera latifolia (2 n  = 4 x  = 60), Chrysolaena flexuosa (2 n  = 2 x  = 20), and Cyrtocymura cincta (2 n  = 4 x  = 40). The relevance of the results is discussed in relation to the available data for each of the analysed taxa. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 221–230.     

Cytotaxonomic analysis of South American species of Vernonia (Vernonieae: Asteraceae)

Chromosome numbers in 48 populations belonging to 31 species of Vernonia from South America have been determined. First chromosome counts are reported for V. propinqua var. canescens (2 n = 20), V. sceptrum (2 n = 80), V. rufogrisea (2 n = 32), V. heringeri (2 n = 32), V. hovaefolia (2 n = 64), V. hystricosa (2 n = 128), V. obtusata (2 n = 64), V. pseudoincana (2 n = 32), V. rubricaulis var.◊ australis (2 n = 64) and V. rugulosa (2 n = 30). The basic chromosome numbers x = 10, 14, 15, 16 and 17 were confirmed for New World species of Vernonia . Together with previous records, these numbers suggest that the evolution of Vernonieae has occurred through a combination of polyploidy and aneuploidy. Taxonomic implications of certain counts are also discussed.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 401–408.