Evolutionary and ecological implications of genome size in the North American endemic sagebrushes and allies (Artemisia, Asteraceae)

The genome size of 51 populations of 20 species of the North American endemic sagebrushes (subgenus Tridentatae ), related species, and some hybrid taxa were assessed by flow cytometry, and were analysed in a phylogenetic framework. Results were similar for most Tridentatae species, with the exception of three taxonomically conflictive species:  Artemisia bigelovii Gray,  Artemisia pygmaea Gray, and  Artemisia rigida Gray. Genome size homogeneity (together with the high morphological, chemical, and karyological affinities, as well as low DNA sequence divergence) could support a recent diversification process in this geographically restricted group, thought to be built upon a reticulate evolutionary framework. The Tridentatae and the other North American endemic Artemisia show a significantly higher genome size compared with the other subgenera. Our comparative analyses including genome size results, together with different kinds of ecological and morphological traits, suggest an evolutionary change in lifestyle strategy linked to genome expansion, in which junk or selfish DNA accumulation might be involved. Conversely, weed or invasive behaviour in Artemisia is coupled with lower genome sizes. Data for both homoploid and polyploid hybrids were also assessed. Genome sizes are close to the expected mean of parental species for homoploid hybrids, but are lower than expected in the allopolyploids, a phenomenon previously documented to be related with polyploidy.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 631–649.     

New or rare chromosome counts in Artemisia L. (Asteraceae, Anthemideae) and related genera from Kazakhstan

Fifteen chromosome counts of six Artemisia taxa and one species of each of the genera Brachanthemum, Hippolytia, Kaschgaria, Lepidolopsis and Turaniphytum are reported from Kazakhstan. Three of them are new reports, two are not consistent with previous counts and the remainder are confirmations of very scarce (one to four) earlier records. All the populations studied have the same basic chromosome number, x = 9, with ploidy levels ranging from 2x to 6x. Some correlations between ploidy level, morphological characters and distribution are noted.     

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.     

Variation of DNA amount in 47 populations of the subtribe Artemisiinae and related taxa (Asteraceae, Anthemideae): karyological, ecological, and systematic implications.

Genome size has been estimated by flow cytometry in 47 populations of 40 species of the tribe Anthemideae (Asteraceae), mainly from Artemisia and other genera of the subtribe Artemisiinae and related taxa. A range of 2C values from 3.54 to 21.22 pg was found. DNA amount per basic chromosome set ranged from 1.77 to 7.70 pg. First genome size estimates are provided for one subtribe, 10 genera, 32 species, and two subspecies. Nuclear DNA amount correlated well with some karyological, physiological and environmental characters, and has been demonstrated as a useful tool in the interpretation of evolutionary relationships within Artemisia and its close relatives.     

CHROMOSOME NUMBERS IN UMBELLIFERAE. III

Chromosome numbers are reported for 156 collections representing 100 taxa of Umbelliferae. Approximately two thirds of the collections are from Mexico, Central and South America and indicate a high percentage of polyploid species in certain genera found in this area. Chromosome numbers for plants belonging to 78 taxa are published here for the first time, previously published chromosome numbers are verified for 18 taxa and chromosome numbers differing from those previously published are reported in seven instances. No chromosome counts have been previously published for nine of the genera included here. Further aneuploidy and polyploidy were found in Eryngium, and Lomatium columbianum has been found to be a high polyploid with 2n = 14x. Every chromosome count is referable to a cited herbarium specimen.     

Cytotaxonomy of Commelinaceae: chromosome numbers of some African and Asiatic species

Chromosome counts are reported for 32 taxa (31 species and 1 subspecies) belonging to 10 genera of Commelinaceae from seven African and Asiatic countries. Counts for 13 species and 1 subspecies are recorded for the first time. Published chromosome numbers for Anhicopsis and Polyspatha are confirmed. It is suggested that Pdisota, Pollia and Stanfieldidla each has a single basic number (x = 20, 16 and 11, respectively). The known cytological diversity in Floscopa is extended. The third continental African species of Coleolrype is found to have the same chromosome number (2n = 36) as the other two. The preponderance of the basic number x = 15 in Commelina is supported. The uncommon basic number x = 13 is reported in four taxa of Cyanotis together with karyotypic differences. The basic number x = 6 is found in a second species of Murdannia . Karyotypic data in addition to chromosome numbers are presented for 24 of the 32 taxa investigated. Karyotypes are found to be useful in assessing relationships in the family, and evolutionary trends in the karyotype are noted.     

CHROMOSOME NUMBERS IN UMBELLIFERAE. IV.

Chromosome numbers are reported for 167 collections representing 100 taxa of Umbelliferae. More than four-fifths of the counts apply to members of subfamilies Hydrocotyloideae (29) and. Saniculoideae (50); the remaining 21 belong to Apioideae. Chromosome numbers of plants belonging to 68 taxa are published here for the first time; chromosome numbers are verified for 23 taxa; and chromosome numbers differing from those published previously are reported in nine instances. No chromosome counts have previously been reported for 19 of the genera included. Polyploidy has been established for Azorella, Mulinum, Coaxana, Enantiophylla, and Tiozimia.     

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.     

Chromosome number variation and polyploidy in the genus Echinocereus (Cactaceae)

Analyses of meiotic and mitotic chromosomes were undertaken in 16 taxa of Echinocereus belonging to 12 species and all seven taxonomic sections (sensu Taylor). Chromosome numbers are reported for the first time for eight taxa, and previously published chromosome counts are confirmed for the remaining eight. Both diploid and polyploid counts were obtained. Eleven (69%) of the taxa surveyed were diploid (2n = 22); the five varieties of E. engelmannii were polyploid (2n = 44). Overall, chromosome counts are available for 23 of the 48 proposed species (sensu Taylor). Of these, 19 (82%) are diploid, and four (18%) are polyploid. Polyploid cytotypes are most common in the primitive sections, e.g., sections Erecti and Triglochidiatus, which suggests that polyploidy is probably a derived condition in Echinocereus. Polyploid taxa range from medium to high latitudes and elevations relative to the overall distribution of the genus. Polyploidy, hybridization, and cryptic chromosomal rearrangements are thought to be the major causes of the speciation events of the genus.     

The Genus Artemisia and its Allies: Phylogeny of the Subtribe Artemisiinae (Asteraceae, Anthemideae) Based on Nucleotide Sequences of Nuclear Ribosomal DNA Internal Transcribed Spacers (ITS)

Abstract: Sequences of the internal transcribed spacers (ITS1 and ITS2) of nuclear ribosomal DNA were analysed for 44 Artemisia species (46 populations) representing all the five classical subgenera and the geographical range of the genus, 11 species from 10 genera closely related to Artemisia, and six outgroup species from five other genera of the Anthemideae. The results definitely support the monophyly of the genus Artemisia in its broadest sense (including some taxa segregated as independent genera, like Oligosporus and Seriphidium ). Eight main clades are established in this molecular phylogeny within Artemisia; they agree in part with the classical subdivision of the genus, but they also suggest that some infrageneric groups must be redefined, especially the subgenus Artemisia. The subgenera Tridentatae and Seriphidium are independent from each other. Some of the satellite genera are clearly placed within Artemisia ( Artemisiastrum, Filifolium, Mausolea, Picrothamnus, Sphaeromeria, Turaniphytum ), whereas some others fall outside the large clade formed by this genus (Brachanthemum, Elachanthemum, Hippolytia, Kaschgaria). Our results, correlated to other data such as pollen morphology, allow us to conclude that the subtribe Artemisiinae as currently defined is a very heterogeneous group. Affinities of the largest genus of the subtribe and tribe, Artemisia, and of other genera of the subtribe to some genera from other subtribes of the Anthemideae strongly suggest that subtribe Artemisiinae needs a deep revision and redefinition. Phylogenetic utility of region trnL-F of the plastid DNA in the genus Artemisia and allies was also evaluated: sequences of the trnL-F region in Artemisia do not provide phylogenetic information.     

Chromosome numbers of Thai Rubiaceae

Chromosome numbers for 14 taxa of indigenous Thai Rubiaceae are presented. They include first counts for 3 genera: Aphaenandra (A. uniflora), Prismatomeris (P. tetrandra subsp. malayana) and Tarennoidea (T. wallichii) ; all show diploidy on x=11. The remaining counts are first counts for species: Argostemma diversifolium, A. neurocalyx and A. pictum, Coptosapelta flavenscens, Gardenia saxtilis, Ixora sp., Morinda sp., Mussaenda sanderiana, Oxyceros horridus, Rothmannia wittii (first count for an Asiatic species of the genus) (all diploid on x=11), and Canthium sp. (tetraploid on x=11). The poor state of karyological knowledge of indigenous Thai Rubiaceae is discussed, and a table including all relevant known chromosome counts is presented. Chromosome data are only known for 38 genera (ca. 41% of all Rubiaceae genera occurring in Thailand); chromosome numbers are often only available for one or few taxa of each genus [in sum, for only about 50 (or for roughly 10% of all) taxa]. Of only 14 genera (ca. 15%), chromosomes were counted from Thai material (for the others, counts originate from elsewhere, i.e. refer to more widely distributed taxa also extending into Thailand).     

Molecular phylogenetic analysis of the Grey-cheeked Fulvetta (Alcippe morrisonia) of China and Indochina: a case of remarkable genetic divergence in a "species"

The Grey-cheeked Fulvetta, Alcippe morrisonia, is a polytypic species of Quaker babbler (Timaliidae) occurring mainly in highlands from Burma across southern China to Taiwan. To examine gene flow among populations, we sequenced the mitochondrial ND2 gene of 39 individuals of six of the seven subspecies, plus multiple individuals of three outgroup Alcippe species. A lack of shared haplotypes and high FST values suggested no gene flow among populations. The nucleotide divergence between geographically juxtaposed subspecies ranged from 0.8% between Guangdong and Hainan to 9.4% between Yunnan and Vietnam. Phylogenetic analysis of the populations yielded a well resolved tree with two major clades. One clade consisted of the geographically central subspecies schaefferi and davidi, which are located largely in the "Mid-central" zoogeographic region of China's "Oriental" realm. The other clade, the geographically peripheral group, consisted of all other A. morrisonia subspecies, as well as an erstwhile outgroup, the Mountain Fulvetta (Alcippe peracensis annamensis) from central Vietnam. This peripheral group was further divided into two clades, one consisting of taxa occurring in China's "Southwest" zoogeographic region (fratercula and A. p. annamensis), and one occurring in China's "Southern" region (morrisonia, rufescentior, and hueti). These three geographic and phylogenetic groups represent at least four different species based on plumage differences and genetic differentiation. The phylogeny provides the first avian molecular evidence of area relationships among China's zoogeographic zones. It also highlights a remarkable and unexpected amount of genetic divergence and structure in a Sino-Indian "species". If such diversity occurs in other groups of birds with similar distribution, the ramifications are important for conservation planning.     

CHROMOSOME NUMBERS IN COMPOSITAE VII: ASTEREAE III

Reports of 129 new chromosome counts are made for the tribe Astereae of Compositae. They are mostly based on determinations of meiotic material, including first counts for one genus and 43 species or subspecies. Counts are now available for more than 63 of the 100-120 genera and 564 of the approximately 2,000 species in the tribe. Three of every four genera with more than one species counted show more than one chromosome number; 15 genera have species with populations with different numbers. Such variation is very high and indicates the need for more detailed cytotaxonomical study in the group.     

Chromosome numbers and karyotype evolution in holoparasitic Orobanche (Orobanchaceae) and related genera

Chromosome numbers and karyotypes of species of Orobanche, Cistanche, and Diphelypaea (Orobanchaceae) were investigated, and 108 chromosome counts of 53 taxa, 19 counted for the first time, are presented with a thorough compilation of previously published data. Additionally, karyotypes of representatives of these genera, including Orobanche sects. Orobanche and Trionychon, are reported. Cistanche (x = 20) has large meta- to submetacentric chromosomes, while those of Diphelypaea (x = 19) are medium-sized submeta- to acrocentrics. Within three analyzed sections of Orobanche, sects. Myzorrhiza (x = 24) and Trionychon (x = 12) possess medium-sized submeta- to acrocentrics, while sect. Orobanche (x = 19) has small, mostly meta- to submetacentric, chromosomes. Polyploidy is unevenly distributed in Orobanche and restricted to a few lineages, e.g., O. sect. Myzorrhiza or Orobanche gracilis and its relatives (sect. Orobanche). The distribution of basic chromosome numbers supports the groups found by molecular phylogenetic analyses: Cistanche has x = 20, the Orobanche-group (Orobanche sect. Orobanche, Diphelypaea) has x = 19, and the Phelipanche-group (Orobanche sects. Gymnocaulis, Myzorrhiza, Trionychon) has x = 12, 24. A model of chromosome number evolution in Orobanche and related genera is presented: from two ancestral base numbers, x(h) = 5 and x(h) = 6, independent polyploidizations led to x = 20 (Cistanche) and (after dysploidization) x = 19 (Orobanche-group) and to x = 12 and x = 24 (Phelipanche-group), respectively.     

New Chromosome Counts of Some Dicots in the Sino-Japanese Region and Their Systematics and Evolutionary Significance

New somatic chromosome numbers for nine species eight families and eight gen era in the Sino-Japanese Region are reported here as shown in Table 1. Data of six genera are previously unknown cytologically. The bearings of these new data on the systematics and evolution of the related species, genera or families are discussed as follows: (1) Platycarya strobilacea Sieb. et Zucc. (Juglandaceae). The chromosome number of this species is 2n=24, with a basic number of x=12, which deviates from 2n=32 occurred in Juglans, Carya, Pterocarya and Engelhardtia with the basic number x= 16. The Juglandaceae appears to be fundamentally paleotetraploid, with an original basic number of x = 6 in Platycarya and x-8 in the other four genera, although secondary polyploidy occurs in Carya. Based on the remarkable morphological differences between Platycarya and the rest seven genera of the family, Manning (1978) established two subfamilies: Platycaryoideae for Platycarya and Juglandoideae for the other genera. Iljinskaya (1990), however, recently established a new subfamily: Engelhardioideae for Engelhardtia. Lu (1982) points out that because of a great number of primitive characters occurring in Platycarya, the genus could not be derived from any other extant juglandaceous taxa but probably originated with the other groups from a common extinct ancestor. The present cytological data gives support to Manning′s treatment. We are also in favor of Lu′s supposition and suggest that basic aneuploid changes, both ascending and descending, from a common ancestor with the original basic number x=7, took place during the course of early evolution of the Juglandaceae and led to the origin of taxa with x=6 and 8. Subsequent polyploidy based on these diploids occurred and brought forth polyploids of relic nature today, whereas their diploid progenitors apparently have become extinct. (2) Nanocnide pilosa Migo (Urticaceae). The chromosome number of this Chinese endemic is 2n-24, with a basic number of x=12. An aneuploid series occurs in the Urticaceae, with x--13, 12, I1, 10, 9, 8, 7, etc. According to Ehrendorfer (1976), x = 14, itself being of tetraploid origin, is the original basic number of the whole Urticales, and descending aneuploid changes took place in the early stage of evolution of the Urticaceae and Cannabinaceae. In addition to Nanocnide, x= 12 also occurs in Australina, Hesperonide and Lecanthus, and partly in Chamabainia, Elatostema, Girardinia, Pouzolzia and Urtica. (3--4) Sedum sarmentosum Bunge and S. angustifolium Z. B. Hu et X. L. Huang (Crassulaceae). The former is a member of the Sino-Japanese Region, while the latter is only confined to eastern China. The chromosome number of Sedum is remarkably complex with n=4-12, 14-16…74, etc. S. angustifolium with 2n=72 of the present report is evidently a polyploid with a basic number of x =18 (9^?) Previous and present counts of S. sarmentosum show infraspecific aneupolyploidy: n = c. 36 (Uhl at al. 1972) and 2n=58 (the present report). These two species are sympatric in eastern China and are morphologically very similar, yet distinguishable from each other (Hsu et al. 1983) S. sarmentosum escaped from cultivation in the United States gardens exhibited high irregularity in meiosis (Uhl et al. 1972). Uhl (pets. comm. ) suspected strongly that it is a highly sterile hybrid. R. T. Clausen (pets. comm.) found that plants of S. sarmentosum naturalized in the American Gardens propagated by means of their long stolons and broken stem tips, and could not yield viable seeds. Hsu et al. (1983) found that some of the plants of S. sarmentosum and S. angustifolium did yield a few seeds, but other did not. These species are, therefore, by the large vegetatively apomictic. (5) Glochidion puberum (L. ) Hutch. (Euphorbiaceae). The genus Glochidion includes about 300 species, but only eigth species from the Himalayas have been studied cytologically, with n= 36 and 2n= 52, having a basic number of x= 13. The present count for the Chinese endemic G. puberum establishes the tetraploid chromosome number 2n= 64, and adds a new basic number x= 16 to the genus. (6) Orixa japonica Thunb. (Rutaceae). Orixa is a disjunct Sino-Japanese monotypic genus. Out of the 158 genera of the Rutaceae, chromosome numbers of 65 genera have hitherto been investigated, of which 42 genera are with x=9 (66.61%), some with x=7, 8 and 10, and rarely with x=13, 15, 17 and 19. The present count of 2n=34 for O. japonica may have resulted from a dibasic tetraploidy of n=8+9. (7) Rhamnella franguloides (Maxim.) Weberb. (Rhamnaceae). The chromosome number of this member of the Sino-Japanese Region is 2n= 24. with a basic number of x= 12. The basic number x= 12 also occurs in Hovenia, Paliurus, Sageretia, Ceanothus and Berchemia. Hong (1990) suggested that x= 12 in Rhamnaceae may be derived from descending aneuploidy of a paleotetraploid ancestor. (8) Sinojackia xylocarpa Hu (Styracaceae). The chromosome number of this rare Chinese endemic is 2n= 24, with a basic number of x =12, which is identical with that in Halesia and Pterostyrax, but deviates from that in Styrax (x=8). The basic number x=8 in the Styracaceae may be derived from the original basic number x=7 by ascending aneuploidy in the early stage of evolution of the family, and x=12 may be derived from polyploidy. (9) Thyrocarpus glochidiatus Maxim. (Boraginaceae). The chromosome number of this Chinese endemic species is 2n=24, with a basic number of x=12. An extensive aneuploid sequence of x = 4-12 occurs in the Boraginaceae, of which x = 8, 7 and 6 are the most common. The basic number x=12 also occurs in Cynoglossum and Mertensia. It is evident that aneuploid changes, both descending and ascending, from an ancestor with x = 7, have taken place in the primary phase of evolutionary diversification of the Boraginaceae, and subsequent polyploidy has given rise to x=15, 17 and 19 in a few genera (e. g. Amsinskia and Heliotropium). The origin of x=12 is not certain. Either it be a result of ascending aneuploidy, or a product of polyploidy on the basis of x = 6. The present authors are in favorof the latter.     

CHROMOSOME COUNTS OF COMPOSITAE FROM THE UNITED STATES AND MEXICO

Chromosome counts are reported for 126 taxa representing 122 species and 61 genera of Compositae. First reports include two genera, Stylocline (n = 14) and Chromolepis (n = 19), 17 species, two infraspecific taxa, and one interspecific hybrid. Five additional taxa have chromosome numbers differing from previously published accounts. Carminatia is reinstated to generic status.     

Intergeneric hybridization between Taeniatherum and different genera of Triticeae, Poaceae

Intergeneric crosses were carried out between the 3 subspecies of Taeniatherum caput-medusae (2x) and 30 different species (33 different cytotypes) representing 11 genera of the Triticeae. Seed set was obtained in 15 combinations, but only 6 of them resulted in adult plants. In the combination with Hordeum bulbosum (2x), a haploid of T. caput-medusae was obtained as a result of selective elimination of the H. bulbosum chromosomes. All other hybrid plants were morphologically intermediate between the parents. They were highly sterile with low pollen fertility caused by very low chromosome pairing at meiosis and without any seed set.
The few hybrid plants obtained and the lack of pairing at meiosis support that Taeniatherum is a distinct genus within the Triticeae.     

CYTOGEOGRAPHIC EVIDENCE FOR THE EVOLUTION OF DISTYLY FROM TRISTYLY IN THE NORTH AMERICAN SPECIES OF OXALIS SECTION IONOXALIS

A survey of haploid chromosome numbers of 18 North American taxa of Oxalis section Ionoxalis was initiated to determine the relationship between ploidal level, geographic distribution, and the occurrence of tristyly and distyly. Although chromosome numbers in the section are variable, the majority of tristylous populations are diploid. Among the distylous taxa a greater diversity of ploidal levels exists, with the higher chromosome numbers predominating. In section Ionoxalis the majority of the tristylous taxa are geographically restricted endemics of southern Mexico, while the distylous taxa have more extensive distributions ranging to the north. The association of diploidy and geographic endemism in the majority of the tristylous taxa suggests that these species are relictual. A few widespread tristylous taxa are polyploid, and often somewhat weedy. The probable derivation of widespread polyploid species from the restricted diploid endemics of southern Mexico appears to have been accompanied by the evolution of distyly from tristyly. The frequent association of polyploidy and distyly in section Ionoxalis has apparently resulted from the concurrence of two evolutionary trends: increase in ploidal level and the derivation of distyly from tristyly.