CHROMOSOME NUMBERS IN SPHAERALCEA SECTION FENDLERIANAE

Chromosome numbers are reported for ten taxa in Sphaeralcea section Fendlerianae (Malvaceae). New ploidy levels are reported for six taxa, with one species not previously reported, and extensive polyploidy at all taxonomic levels is documented. The geographic and taxonomic distribution of polyploids suggests that polyploidy arose many times in the taxa of Sphaeralcea. Sphaeralcea fendleri var. venusta, S. polychroma, and S. wrightii populations have yielded exclusively tetraploid counts. Tetraploidy is correlated with taxa having lavender petals. Polyploidy has also allowed the taxa to expand their distributions without resulting in speciation.     

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.     

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 THE COMPOSITAE. XIV. LACTUCEAE

Reports of 150 original chromosome counts are recorded, including reports of 22 genera and 57 species and subspecific taxa in tribe Lactuceae. Also included are first reports for 12 specific or subspecific taxa. x = 9 appears to be the ancestral base of the tribe. Chromosome numbers are known for over 85% of the genera of the tribe and the frequency of polyploidy is ca. 23%, which is about one-half that of the angiosperms.     

Cytotaxonomy and breeding system of the genusBiscutella (Cruciferae)

Chromosome counts were determined for 46 populations ofBiscutella representing 28 taxa. The genus was found to contain diploid taxa with 2n = 12, 16 and 18, tetraploid taxa with 2n = 36 and hexaploid taxa having 2n = 54.B. laevigata L. s. l. consists of diploid and tetraploid populations which are poorly differentiated morphologically. TetraploidB. laevigata s. l. and hexaploidB. variegata Boiss. & Reuter (s. l.) are characterized by chromosomal instability. The variation in chromosome numbers and the occurrence of polyploidy is discussed in relation to the taxonomy of the genus. An investigation of the breeding system showed that most of the annual species were self-compatible and partly inbreeding and most of the perennial species self-incompatible and, therefore, outbreeding, while one annual species,B. cichoriifolia Loisel., showed both systems.     

CHROMOSOME NUMBERS IN THE GENUS ANTHURIUM (ARACEAE) II

Chromosome numbers were determined for 86 Anthurium species. Fifty-one of these were newly determined with counts ranging from 2n = 24 to 66 and 30 being the most common. All known Anthurium chromosome numbers were summarized, and 43 taxonomic changes were made in the previous reports to reflect current taxonomy. In terms of somatic chromosome numbers, the numbers form four polyploid series of 20–40–60, 24–30–48–84, 28–56 and 30–60–90–ca. 124. Paleoaneuploidy, polyploidy and B-chromosomes are basic features of the genus, but subsequent recent aneuploidy is not. The exact nature of chromosome evolution in Anthurium remains to be elucidated.     

Karyotype and genome size variation in Ajuga L. (Ajugoideae–Lamiaceae)

Chromosome number changes and karyotype evolution play an important role in plant genome diversification and eventually in speciation. The genus Ajuga L. (Lamiaceae) has approximately 50 species distributed in temperate to subtropical regions. Four of these species are currently recognized in Korea (A. decumbens Thunb., A. multiflora Bunge, A. nipponensis Makino and A. spectabilis Nakai). Understanding the karyotype evolution in Ajuga has been hampered by the small size of their chromosomes and symmetrical karyotypes. Here we used classic Feulgen staining to establish chromosome numbers and construct karyotypes of the four species of Ajuga recognized in Korea and flow cytometry was used to study their variation in genome. The chromosome number of all investigated plants was 2n = 32. Still, the 2C DNA content ranged from 2.18 pg (A. decumbens) to 4.53 pg (A. multiflora). While the chromosome numbers were the same for all investigated species, the genome size variation could potentially be used as a taxonomic marker.     

CHROMOSOME COUNTS AND TAXONOMIC NOTES FOR COMPOSITAE FROM THE UNITED STATES AND MEXICO

Chromosome counts are reported for 167 taxa representing 160 species and 76 genera of Compositae. First reports for 27 species and 2 infraspecific taxa are included. New chromosome numbers are reported for 9 additional taxa. The genus Carminatia DC. is merged with Brickellia Ell. resulting in two new combinations: Brickellia tenuiflora (DC.) Keil & Pinkava and Brickellia recondita (McVaugh) Keil & Pinkava.     

ISOZYME NUMBER IN SUBTRIBE ONCIDIINAE (ORCHIDACEAE): AN EVALUATION OF POLYPLOIDY

The Oncidiinae has attracted attention because of the variation it exhibits in chromosome number, n = 5–30, which is greater than the range in the rest of the Orchidaceae. The genus Psygmorchis, with n = 5 and 7, has been a particular focus of controversy, and many authors have suggested that 5 and 7 are the base numbers for the subtribe. The other taxa in the subtribe presumably evolved through hybridization and polyploidy. Other workers have found that the lowest counts correlate with derived morphological conditions and have hypothesized that these low numbers result from aneuploid reductions, while higher numbers are associated with ancestral morphologies and are not the result of polyploidy. These two hypotheses were evaluated by determining isozyme numbers for 13 enzymes in species that span the chromosomal range known for the Oncidiinae (n = 5–30). Isozyme number has been shown to be a reliable indicator of polyploidy in angiosperms because polyploids display isozyme multiplicity relative to diploids. This analysis revealed no differences among species in isozyme number for the enzymes examined. Therefore, our data reject the hypothesis that species with higher chromosome numbers are polyploid.     

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.     

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.     

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.     

CHROMOSOME NUMBERS AND TAXONOMIC NOTES FOR ROCKY MOUNTAIN UMBELLIFERAE

Chromosome numbers are reported for 111 collections of Umbelliferae, which represent 38 species belonging to 23 genera. Three of these genera, Aletes, Neoparrya, and Musineon, have not been counted previously. In addition, first counts are presented for Angelica grayi, A, roseana, Cymopterus acaulis, C. bulbosus, C. longipes, C. montanus, Lomatium foeniculaceum subsp. macdougalii, L. megarrhizum, L. nuttallii, L. orientate, and L. simplex. Our reports of polyploidy in Ligusticum ported (hexaploid, 2n = ca. 66) and in Pteryxia terebinthina var. calcarea (tetraploid, 2n = 44) are the first for these taxa. The counts of 2n = 22 for Harbouria trachypleura differ from the previous report of 2n = 20.     

Chromosome numbers in Heterotheca,including Chrysopsis (Compositae: Astereae), with Phylogenetic interpretations

A summation of both previously reported and original data on chromosome numbers is presented for species ofHeterotheca sens. lat. Chromosome numbers are listed for 24 taxa, including 20 different species. Chromosome numbers for the genus aren = 4, 5, 9, 12, and 18. The most common and possibly basic number for the genus isn = 9, with then = 18 taxa considered to be tetraploids, and those withn = 4 and 5 as probable aneuploid derivatives ; alternativelyn = 4 and 5 could be considered the basic numbers withn = 9 the result of polyploidy. Several nomenclatural changes have been made including the following new combinations:Heterotheca bolanderi (Gray) Harms,H. fastigiata (Greene) Harms, andH. villosa var.hispida (Hook.) Harms.     

PHYLOGENETIC IMPLICATIONS OF DIVERSE SEED TYPES,CHROMOSOME NUMBERS,AND POLLEN MORPHOLOGY IN HOUSTONIA (RUBIACEAE)

Seed and pollen morphology were studied by light microscopy and scanning electron microscopy in 39 North and Central American species of Houstonia (including Hedyotis, but excluding Oldenlandia). Chromosome counts were obtained for eight taxa, of which five lacked previous chromosome data. A chromosome number of n = 17 for Houstonia gracilis is a new base number for the genus. Seed external morphology in the genus is very diverse, including variation in compression, margins, testa surfaces, and elaboration of ventral cavities or depressions and hilar ridges or their absence. Three types of pollen apertures are recognized: colporate with type A os, colpororate, and colporate with type B os, the last the most advanced type, occurring in H. caerulea and related species. The 39 species are arranged in twelve groups, based on correlation of seed, pollen, and chromosome data. Geographic distribution provides supplementary evidence for the distinctness and integrity of the six principal groups each composed of 2–9 species. Five of the six minor groups each with one species need chromosome data to facilitate future taxonomic decisions. Chromosome numbers of x = 6, 7, 8, 9, 10, 11, 13, and 17 are now known in this genus, and phylogenetic implications of the combined data are discussed.     

Molecular phylogenetic analyses of nuclear and plastid DNA sequences support dysploid and polyploid chromosome number changes and reticulate evolution in the diversification of Melampodium (Millerieae, Asteraceae)

Chromosome evolution (including polyploidy, dysploidy, and structural changes) as well as hybridization and introgression are recognized as important aspects in plant speciation. A suitable group for investigating the evolutionary role of chromosome number changes and reticulation is the medium-sized genus Melampodium (Millerieae, Asteraceae), which contains several chromosome base numbers (x = 9, 10, 11, 12, 14) and a number of polyploid species, including putative allopolyploids. A molecular phylogenetic analysis employing both nuclear (ITS) and plastid (matK) DNA sequences, and including all species of the genus, suggests that chromosome base numbers are predictive of evolutionary lineages within Melampodium. Dysploidy, therefore, has clearly been important during evolution of the group. Reticulate evolution is evident with allopolyploids, which prevail over autopolyploids and several of which are confirmed here for the first time, and also (but less often) on the diploid level. Within sect. Melampodium, the complex pattern of bifurcating phylogenetic structure among diploid taxa overlain by reticulate relationships from allopolyploids has non-trivial implications for intrasectional classification.     

CHROMOSOME NUMBERS IN THE PHASEOLEAE (FABACEAE:FABOIDEAE) AND THEIR RELATION TO TAXONOMY

Chromosome numbers are reported for 33 species of the tribe Phaseoleae. Six reports are first counts for their species; one report (Ophrestia hedysaroides) is a first count for the genus. This increases the number of genera counted to 53 out of a total 84 for the tribe. A survey of base numbers shows a general pattern of numbers 10 or 11, the same base numbers as in the neighboring tribes Dalbergieae sensu lato, Galegeae sensu lato, and Abreae. The chromosomes are generally small and polyploidy is uncommon. Deviations from base numbers 10 or 11 are mostly found in those genera with morphological pecularities and puzzling taxonomic placements: Erythrina (21), Clitoria (8, 11, 12), Butea (9), Calopogonium (18), Teramnus (14), and Strongylodon (14). Two genera have base numbers which suggest derivation by polyploidy: Glycine (20) and Cologania (22).     

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.     

CYTOLOGICAL STUDIES IN THE GENUS DESMODIUM; SOME CHROMOSOME COUNTS

Chromosome counts for 30 collections representing 21 species examined by the authors are reported. Of these, eight species counts are a first report; the others are confirmation of earlier reports. The genus Desmodium is shown to be dibasic with base chromosome numbers of x = 10 and x = 11.     

ON THE METHODS OF EXPERIMENTAL TAXONOMY

By studying seedling progenies from individual plants it may be decided whether the material under investigation is allogamous, autogamous or apomictic. Chromosome counts disclose whether the material is cytologically stable or variable. If variation occurs it may be a question of polyploidy or aneuploidy. Aneuploidy may either be an occasional consequence of polyploidy or it may represent dysploidy and the formation of new basic numbers. Chromosome counts combined with measurements of chromosome size may reveal cases of pseudopolyploidy of the kind first observed in Luzula. If chromosome morphology is different between two taxa this indicates reproductive isolation. Different species usually differ with regard to their chromosome structure. Studies of meiosis are desirable in polyploids in order to distinguish between auto- and alloploidy. Cases of genic control of meiosis in polyploids of Phleum and Triticum demonstrate, however, that partial or complete homology between genomes does not always result in multivalent formation. Apparent alloploids may in reality be more or less autoploid. Incompatibility is not a reliable criterion of species differentiation. Strong or absolute barriers between diploids and autotetraploids may have purely quantitative causes. Hybrid sterility is not always a reliable measure of the degree of genetic difference between the parents, but may be caused by heterozygosity for chromosomal rearrangements. Cytological and experimental methods are, nevertheless, indispensable tools in plant taxonomy.