CHROMOSOME NUMBERS OF CAMPANULACEAE. III. REVIEW AND INTEGRATION OF DATA FOR SUBFAMILY LOBELIOIDEAE

Chromosome numbers are now known for 153 species in 21 genera of Lobelioideae (Campanulaceae); this represents almost 13% of the species and 70% of the genera in the subfamily. Numbers reported are n = 6, 7, 8, 9, 10, 11, 12, 13, 14, 19, 21, 35, 70. The subfamily as a whole has x = 7; the best documented exception is Downingia and its allies with x = 11. Only four genera show interspecific variation in chromosome number: Downingia (n = 6, 8, 9, 10, 11, 12); Lobelia (n = 6, 7, 9, 12, 13, 14, 19, 21); Pralia (n = 6, 7, 13, 14, 21, 35, 70); and Solenopsis (n = 11, 14). Intraspecific variation occurs in 13 species, with as many as four different cytotypes in one species. The herbaceous members of the subfamily as a group are quite variable, showing the entire range of chromosome numbers, including numerous dysploids, but are predominantly diploid. The woody species, by contrast, are much less variable; nearly all of the species are tetraploid, with only a few diploids and hexaploids and no dysploid numbers known. These data support the hypothesis that woodiness is apomorphic within the subfamily. A general trend of higher chromosome numbers at higher latitudes and higher elevations is evident within the subfamily. The chromosome number of Apetahia raiateensis (n = 14) is reported here for the first time, on the basis of a count made about 30 years ago by Peter Raven.     

Chromosome numbers of miscellaneous Malagasy Acanthaceae

Meiotic chromosome numbers are reported for 12 species in eight genera of Acanthaceae from Madagascar. Chromosome numbers of 11 species are reported for the first time. Counts inMendoncia (n=19) andNeuracanthus (n=20) are the first for these genera. A new chromosome number (n=30) is reported inJusticia. Systematic implications of the chromosome counts are addressed and basic chromosome numbers for these eight genera of Malagasy Acanthaceae are discussed.     

CHROMOSOME NUMBERS OF HAWAIIAN FLOWERING PLANTS AND THE SIGNIFICANCE OF CYTOLOGY IN SELECTED TAXA

Chromosome numbers are reported for 128 species of flowering plants indigenous or endemic to Hawaii, including first reports for 13 genera and 82 species. The special significance of reports for Ilex, Tribulus, Keysseria, Pisonia, Boerhavia, Jacquemontia, Claoxylon, Lipochaeta, Railliardia, and Dubautia are discussed. The cytological and morphological variation in Railliardia and Dubautia is considered and their treatment as congeners is advocated. The cytogeogruphic pattern in Dubautia and Railliardia and other factors suggest that the ancestral chromosome number of the Hawaiian tarweeds is n = 14. Their derivation from a western North American progenitor similar to Adenothamnus is considered plausible if not indeed likely.     

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.     

CHROMOSOME NUMBERS IN CUPHEA (LYTHRACEAE): NEW COUNTS AND A SUMMARY

The American genus Cuphea with ca. 260 species is extremely diverse with respect to chromosome number. Counts are now available for 78 species and/or varieties, or 29% of the genus. Included in this study are first reports for 15 taxa from Brazil, Cuba, Dominican Republic, Mexico, and Venezuela. Twenty-two different numbers are known for the genus, ranging from n = 6 to n = 54. The most common number in the primary center of species diversity in Brazil is n = 8, which is regarded as the base number of the genus. Two numbers are most common in the secondary center in Mexico, n = 10 and n = 12. Species with n = 14 or higher are considered to be of polyploid origin. Polyploids comprise 46% of the total species counted and appear in 9 of the 11 sections for which chromosome numbers have been reported. Aneuploid species comprise ca. 25% of the genus and are known from 7 of the 11 sections. The two subgenera are not characterized by different chromosome numbers or sequences of numbers. None of the 14 sections are circumscribed by a single chromosome number. Morphological and ecological variability in widespread, weedy species is correlated with differing chromosome numbers in some species whereas in others the chromosome number is stable. Summary of chromosome numbers by taxonomic section is presented. Section Euandra, centered in eastern Brazil, and the largest section of the genus, appears to be chromosomally most diverse. In section Trispermum, characterized by difficult, variable species with intermediate forms, two of the four species studied have polyploid races. Section Heterodon, endemic to Mexico and Central America and comprising most of the annual species of the genus, is best known chromosomally. Chromosome numbers have been counted for 25 of 28 species, and 12 different numbers are reported. The most advanced sections, Melvilla and Diploptychia, with numerous species occurring at higher altitudes, are characterized by high polyploids. Apomictic species occur in sect. Diploptycia. The cytoevolution of Cuphea is complex with frequent polyploid and aneuploid events apparently playing a significant role in speciation in both centers of diversity.     

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.     

CHROMOSOME NUMBERS IN THE COMPOSITAE: COLOMBIAN SPECIES

Chromosome numbers are presented for 76 species belonging to 35 genera of Compositae from Colombia. Thirty-nine species and three genera, Espeletia (x = 19), Steiractinia (x = 14), and Vasquezia (x = 19), are reported for the first time. New base numbers or chromosome series are recorded in Baccharis (B. nitida, n = 25), Calea (C. caracasana, n = 24), and Liabum (L. mega-cephalum, n = 10).     

CHROMOSOME NUMBERS OF CAMPANULACEAE. II. THE LOBELIA TUPA COMPLEX OF CHILE

Gametic chromosome numbers are reported for 27 collections representing the four species of the Lobelia tupa complex (Campanulaceae, Lobelioideae) in Chile; all are n = 21. This represents the first report of chromosome numbers for L. bridgesii Hook. & Arn., L. excelsa Bonpl., and L. polyphylla Hook. & Arn., and confirms previous reports of this number in L. tupa L. As the basic chromosome number of Lobelioideae is x = 7, these species are interpreted as hexaploids. Higher polyploids are extremely rare among Lobelioideae; most of those previously reported have been either sporadic individuals or populations within an otherwise diploid or tetraploid species, or occasional species within an otherwise diploid and tetraploid lineage. This is the first report of an entire complex of lobelioid species that is uniformly hexaploid. This suggests that the Chilean endemics are relatively derived within Lobelia, and offers some support for the monophyly of the complex.     

Karyological features and cytotaxonomy of the tribe Vernonieae (Asteraceae)

Chromosome numbers are reported for 29 populations of 19 Vernonieae taxa collected mainly in the northeastern region of Brazil. Among them, data for five genera (Blanchetia, Rolandra, Pithecoseris, Stilpnopappus and Vanillosmopsis) are here reported for the first time, and the first chromosome counts are presented for 12 species. Chromosome numbers are quite diverse among and sometimes within genera, especially in the controversial and large subtribe Vernoniinae. The numbers varied from 2n = 18 to 2n = ~72. The main karyoevolutionary mechanism seems to be dysploidy, while polyploidy is probably associated with ancient hybridization processes generating most paleotetraploid genera. All studied species presented semi-reticulated interphase nuclei and proximal-early condensing behavior in prophase to prometaphase. In one species (Vernonia condensata with 2n = 40) fluorochrome staining with CMA/DAPI revealed five chromosome pairs bearing subterminal CMA⁺/DAPI^? heterochromatin, probably NOR-associated, revealing the existence of low amounts of satellite DNA. The role of these features in the evolution of the tribe is discussed, revealing some interesting aspects for understanding of the Vernonieae karyoevolution, especially regarding neotropical members.     

CHROMOSOME NUMBERS IN COMPOSITAE. III. SENECIONEAE

Ornduff , Robert (Duke U., Durham, N. C), Peter H. Raven , Donald W. Kyhos , and A. R. Kruckeberg . Chromosome numbers in Compositae. III. Senecioneae. Amer. Jour. Bot. 50(2): 131–139. Illus. 1963.—Chromosome counts are reported for 75 taxa of tribe Senecioneae (Compcsitae) and are listed with a generic summary of previous counts in the tribe. First counts are reported for Bedfordia, Crocidium, Dimeresia, Gamolepis, Lepidospartum, Luina, Peucephyllum, Telradymia, and the first definite count recorded for Euryops. New numbers are added to those previously known in Arnica and Psathyrotes. Intraspecific differences in ploidy-level are reported in 4 North American species of Senecio. Although chromosome numbers are useful as an aid in delimiting some genera of Senecioneae, they are of little use in circumscribing genera peripheral to Senecio, primarily because of the great range of chromosome numbers of that genus. Chromosome numbers support suggestions based on morphological considerations that genera such as Crocidium and Dimeresia do not belong in Senecioneae, whereas chromosome number and morphology of the plants virtually prohibit the removal of such genera as Peucephyllum, Lepidospartum, and Telradymia from Senecioneae, despite the suggestions of several recent authors. It is proposed that the base number for the tribe is 10 and that the tribe originated in the Old World, with subsequent widespread migration and diversification.     

CHROMOSOME NUMBERS IN COMPOSITAE. VI. SENECIONEAE. II

Chromosome counts are reported for 76 taxa and 2 natural hybrids of tribe Senecioneae (Compositae). First counts are reported for several species of Senecioneae as well as for the genera Cadiscus and Whitneya. New chromosome numbers are added to those previously known in Arnica, Cacalia, and Senecio. Additional counts from Arnica support our previous suggestion that x = 19 for this genus. It is assumed that observed meiotic irregularities are associated with apomixis in this genus. Basic chromosome numbers for various New World sections of Senecio are proposed, and certain problems of sectional relationships in this genus are discussed. Chromosome numbers and plant morphology of Cadiscus, Hulsea, and Whitneya indicate that these genera should be removed from Helenieae to Senecioneae. The possible affinity of the anomalous genus Adenocaulon with Mutisieae is discussed. Data presented in the paper further support our earlier proposal that the basic chromosome number for Senecioneae is x = 10.     

Interstitial telomere‐like repeats in the monocot family Araceae

Combining molecular cytogenetics and phylogenetic modelling of chromosome number change can shed light on the types of evolutionary changes that may explain the haploid numbers observed today. Applied to the monocot family Araceae, with chromosome numbers of 2n = 8 to 2n = 160, this type of approach has suggested that descending dysploidy has played a larger role than polyploidy in the evolution of the current chromosome numbers. To test this, we carried out molecular cytogenetic analyses in 14 species from 11 genera, using probes for telomere repeats, 5S rDNA and 45S rDNA and a plastid phylogenetic tree covering the 118 genera of the family, many with multiple species. We obtained new chromosome counts for six species, modelled chromosome number evolution using all available counts for the family and carried out fluorescence in situ hybridization with three probes (5S rDNA, 45S rDNA and Arabidopsis‐like telomeres) on 14 species with 2n = 14 to 2n = 60. The ancestral state reconstruction provides support for a large role of descending dysploidy in Araceae, and interstitial telomere repeats (ITRs) were detected in Anthurium leuconerum, A. wendlingeri and Spathyphyllum tenerum, all with 2n = 30. The number of ITR signals in Anthurium (up to 12) is the highest so far reported in angiosperms, and the large repeats located in the pericentromeric regions of A. wendlingeri are of a type previously reported only from the gymnosperms Cycas and Pinus. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 15–26.     

CHROMOSOME NUMBERS IN SOUTH AMERICAN HAPLOPAPPUS CASS. (COMPOSITAE)

Chromosome counts are reported for 33 species from all four sections of the genus Haplopappus in South America. These include first reports for 28 species and two putative hybrids. All chromosome numbers reported herein are 2n = 5_II, with the exception of H. prunelloides with 2_n = 6_II. Unlike the North American species, the morphological diversity of South American taxa is not concomitant with chromosomal variation.     

Karyological studies on some Antarctic liverworts

Abstract

Chromosome counts are reported from five species of Antarctic hepatics belonging to five genera and four families. The chromosome numbers in the genera Herzogobryum (H. teres n = 9) and Paehyglossa (P. dissitifolia n = 9) are reported for the first time. The first chromosome count from Lophozia excisa, n = 27, is the highest reported from Lophozia. A diploid number, n = 18, is reported for the first time from Barbilophozia hateheri; a second count, n = 9, agrees with earlier reports. The commonest Antarctic liverwort Cephaloziella cf. exili[lora has n = 18 and this agrees with earlier reports for the genus.     

CHROMOSOME NUMBERS IN PERITYLE AND RELATED GENERA (PERITYLANAE-COMPOSITAE)

Chromosome numbers are presented for 28 species of the genus Perityle, one putative inter-sectional hybrid, two species of Amauria, one species of Eutetras, and one species of Pericome. For Perityle, initial counts are recorded for 12 species of sect. Laphamia (n = 16, 17, 18, 36, ca. 102) and 11 species of sect. Perityle (n = 11, 12, 13, 16, 17, 18, 19, 34, 51). Chromosome numbers for the two species of Amauria (n = 18) are first reports for the genus. Including the current information, chromosome numbers have been recorded for 37 of the approximately 50 species recognized for Perityle. At least 24 taxa have numbers of n = 17, suggesting a base chromosome number of x = 17 for Perityle.     

Karyological knowledge of the Italian vascular flora as inferred by the analysis of “Chrobase.it”

Abstract

Chromosome number knowledge of the Italian vascular flora is stored in the online database Chrobase.it, which includes 6723 records, referable to 3428 taxa, 2799 accepted species and subspecies (about 35% of the national flora), and 3410 different chromosome countings (cytotypes). Appropriate queries to Chrobase.it allowed us to calculate mean, modal and median chromosome numbers for the Italian vascular flora, for geographical subgroups (islands, south, centre, north) and for selected orders, families and genera. Chromosome number data were available for 41 out of 55 orders (74%) and 107 out of 428 families (67%), represented by 664 out of 1297 genera (51%). The most studied administrative regions are Sicily (844 taxa), Tuscany (592 taxa), and Sardinia (390 taxa), while the most studied families are Asteraceae (465 taxa), Fabaceae (266 taxa), Brassicaceae (158 taxa), and Poaceae (144 taxa). Chromosome numbers range from 2n = 6, occurring in several species of Hypochaeris (Asteraceae), to 2n = 240, occurring in Ophioglossum (Ophioglossaceae), Dryopteris (Dryopteridaceae) and Arenaria (Caryophyllaceae) (mode is 2n = 18, and median is 2n = 24). Chromosome number variability was analyzed by frequencies (linear plots) and ANOVA, resulting in significant differences among geographical groups (mean chromosome number increasing from islands-south to centre-north) and selected taxa. B-chromosomes occur in 5.3% of data (148 taxa) and their number is not significantly different among geographical areas, while they occur only in 14 orders, 17 families, and 56 genera. The number of B-chromosomes ranges from 1 to 13 (mode = 1, median = 2).     

Chromosome counts and karyotypes in Chaetoseris and Stenoseris (Asteraceae‐Cichorieae) from the Hengduan Mountains of SW China

Abstract Chaetoseris and Stenoseris are two morphologically close genera from the tribe Cichorieae of the sunflower family and they are endemic in alpine eastern Himalayas to the Hengduan Mountains of SW China. Mitotic chromosome numbers and karyotypes are reported for 12 populations representing eight species of Chaetoseris and two species of Stenoseris from the Hengduan Mountains region. Eight species are new and the other two provide confirmation of previous reference. All Chaetoseris and Stenoseris taxa are diploidy with 2n= 16 and their basic number is tentatively suggested as x= 8. Karyotypes of Chaetoseris and Stenoseris are similar to each other with 2A and 2B for the former and 2A for the latter. Cytological data of chromosomal numbers and karyotypes support a close relationship of the two genera. Currently no polyploids are found for these two genera and it seems that polyploidization has played a minor role in their evolutionary speciation in the Hengduan Mountains region.     

New chromosome counts and evidence of polyploidy in Haageocereus and related genera in tribe Trichocereeae and other tribes of Cactaceae

Chromosome numbers for a total of 54 individuals representing 13 genera and 40 species of Cactaceae, mostly in tribe Trichocereeae, are reported. Five additional taxa examined belong to subfamily Opuntioideae and other tribes of Cactoideae (Browningieae, Pachycereeae, Notocacteae, and Cereeae). Among Trichocereeae, counts for 35 taxa in eight genera are reported, with half of these (17 species) for the genus Haageocereus. These are the first chromosome numbers reported for 36 of the 40 taxa examined, as well as the first counts for the genus Haageocereus. Both diploid and polyploid counts were obtained. Twenty nine species were diploid with 2n=2x=22. Polyploid counts were obtained from the genera Espostoa, Cleistocactus, Haageocereus, and Weberbauerocereus; we detected one triploid (2n=3x=33), nine tetraploids (2n=4x=44), one hexaploid (2n=6x=66), and three octoploids (2n=8x=88). In two cases, different counts were recorded for different individuals of the same species (Espostoa lanata, with 2n=22, 44, and 66; and Weberbauerocereus rauhii, with 2n=44 and 88). These are the first reported polyploid counts for Haageocereus, Cleistocactus, and Espostoa. Our counts support the hypothesis that polyploidy and hybridization have played prominent roles in the evolution of Haageocereus, Weberbauerocereus, and other Trichocereeae.     

Chromosome numbers and generic relationships in subtribe Stephanomeriinae (Compositae: Ichorieae)

Chromosome numbers are reported from over 230 populations representing species in eight genera. First counts are reported for three species ofStephanomeria, five species ofLygodesmia, and one species ofPinaropappus. Base chromosome numbers,x = 6, 7, 8, and 9 are known in the subtribe;x = 9 is found in six of the 12 genera and presumably is the ancestral base number for the subtribe. Two phyletic lines, aMalacothrix line and aStephanomeria line are recognized on morphological grounds. A key to the 12 genera is provided.     

Evolutionary trends in Iridaceae: new cytogenetic findings from the New World

With the present work, we aim to provide a better understanding of chromosome evolutionary trends among southern Brazilian species of Iridoideae. Chromosome numbers and genome sizes were determined for 21 and 22 species belonging to eight genera of Tigridieae and two genera of Trimezieae, respectively. The chromosome numbers of nine species belonging to five genera are reported here for the first time. Analyses of meiotic behaviour, tetrad normality and pollen viability in 14 species revealed regular meiosis and high meiotic indexes and pollen viability (> 90%). The chromosome data obtained here and compiled from the literature were plotted onto a phylogenetic framework to identify major events of chromosome rearrangements across the phylogenetic tree of Iridoideae. Following this approach, we propose that the ancestral base chromosome number for Iridoideae is x = 8 and that polyploidy and dysploidy events have occurred throughout evolution. Despite the variation in chromosome numbers observed in Tigridieae and Trimezieae, for these two tribes our data provide support for an ancestral base number of x = 7, largely conserved in Tigridieae, but a polyploidy event may have occurred prior to the diversification of Trimezieae, giving rise to a base number of x₂ = 14 (detected by maximum‐parsimony using haploid number and maximum likelihood). In Tigridieae, polyploid cytotypes were commonly observed (2x, 4x, 6x and 8x), whereas in Trimezieae, dysploidy seems to have been the most important event. This feature is reflected in the genome size, which varied greatly among species of Iridoideae, 4.2‐fold in Tigridieae and 1.5‐fold in Trimezieae. Although no clear difference was observed among the genome sizes of Tigridieae and Trimezieae, an important distinction was observed between these two tribes and Sisyrinchieae, with the latter possessing the smallest genome sizes in Iridoideae. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 177 , 27–49.