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.     

A cytological study ofPelargonium sect.Eumorpha (Geraniaceae)

The chromosome numbers of seven species ofPelargonium sect.Eumorpha have been determined from material of known wild origin, and karyotypic comparisons have been made. Within the section there is variation in basic chromosome number (x = 4, 8, 9, 11), variation in chromosome size, and two species have polyploid races. The three species with chromosome numbers based on x = 11 have the smallest chromosomes (1.0–1.5 µm); chromosomes are larger (1.0–3.0 µm) in the other species.P. elongatum has the lowest chromosome number in the genus (2n = 8).P. alchemilloides is exceptional in that it has four cytotypes, 2n = 16, 18, 34 and 36, and the form with 2n = 36 has large chromosomes (2.0–5.0 µm). Evidence from a synthesized hybrid suggests thatP. alchemilloides with 2n = 16 may be of polyploid origin. The three species based on x = 11 appear to be more closely related to species from other sections ofPelargonium that have the same basic chromosome number and small chromosome size, rather than to other species of sect.Eumorpha.     

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.     

Chloroplast DNA restriction site data support a narrowed interpretation ofEupatorium (Asteraceae)

Chloroplast DNA (cpDNA) restriction site analysis was used to assess relationships among samples of Eupatorieae from eastern North America. A total of 270 cpDNA variants was recorded from 35 species using 13 restriction enzymes. Phylogenetic analysis usingGalinsoga, Flaveria, andHelianthinae as outgroups indicated that samples ofAgeratina, Hofmeisteria, Mikania, andStevia, each of which have relatively high base chromosome numbers, formed an unresolved basal polytomy. The remaining samples examined formed a well-supported clade, within which there was a split betweenBrickellia, which hasx = 9, and a number of genera withx = 10. Within thex = 10 clade, there was an unresolved trichotomy, with one group ofAgeratum, Conoclinium, Fleischmannia, andCritonia, a second ofLiatris andTrilisa, and a third ofEupatorium s.s. Within theEupatorium s.s. clade there were three further clades, withE. sect.Verticillata diverging first, and a subsequent split between species originating from North America and those from Asia. The cpDNA restriction site data provided support for a relatively narrow interpretation ofEupatorium, and indicated that a high chromosome base number is plesiomorphic for Eupatorieae.     

CHROMOSOMES AND SYSTEMATICS OF ASPLENIUM SECT. HYMENASPLENIUM (ASPLENIACEAE)

Gametic and somatic chromosome numbers of eight Asian species belonging to Asplenium sect. Hymenasplenium were determined. Seven species were observed to have chromosome numbers based on x = 39, and one on x = 38. These chromosome numbers are exceptional in Asplenium which is well known to have chromosomes of n = 36 or multiples thereof. Decisions on the taxonomic status of the species of section Hymenasplenium were facilitated by cytological observations. Systematic recognition of this section is also supported by the peculiarity in the chromosome numbers, and evidence for the addition to the section of several other species is presented.     

Chromosome numbers of Panamanian Lecythidaceae and their use in subfamilial classification

Nine species of Lecythidaceae subfamily Lecythidoideae in four genera whose chromosome numbers were previously unknown, have 17 as their basic chromosome number:Eschweilera pittieri, three other unidentified species ofEschweilera, Grias cauliflora, Gustavia dubia, G. superba, Lecythis minor, andL. tuyrana. All are diploid exceptGustavia superba, which is tetraploid.Couroupita guianensis, which was previously—and probably incorrectly—reported to have a gametic chromosome number of 18, also hasn = 17. The known chromosome numbers support recognizing at least three of Niedenzu’s subfamilies: Planchonioideae withx = 13, Napoleonaeoideae withx = 16, and Lecythidoideae withx = 17. His fourth subfamily, Foetidioideae, with one genus of five species, has not been counted. Cytological data have been and probably will be useful in indicating to what subfamily problematic genera belong and in showing interesting phytogeographic patterns within the family. On the other hand, cytological data provide no recognizable clues relating the Lecythidaceae to other families.     

Die Chromosomenzahlen der in Mitteleuropa vorkommenden Arten vonOrobanche sect.Orobanche

The chromosome numbers of five species ofOrobanche sect.Orobanche (O. alsatica, O. laserpitii-sileris, O. loricata, O. salviae, O. teucrii) are reported for the first time and previous counts could be verified in ten other species. Now the chromosome numbers of all species of sect.Orobanche occurring in Central Europe are known: they are diploid (2n = 38) with the exception ofO. gracilis (tetra- and hexaploid, aneusomatic).

     

A classification of the New World species of Aster (Asteraceae)

In order effectively to list and comment on the results of cytological investigations inAster in a companion paper, a scheme of infrageneric classification is presented which utilizes the basic chromosome number as a pivotal diagnostic character. Reasons are stated as to why, with the exception ofUnamia Greene which is transferred toSolidago, and the commonly recognized generaLeucelene Greene,Machaeranthera Nees andXylorhiza Nutt., none of the segregate genera previously proposed or recorded in the literature is upheld. Instead, these taxa are being given subgeneric or sectional rank. Two additional subgenera are established to accommodate the species groups traditionally placed in “Aster proper,” which are characterized by having basic chromosome numbers ofx = 5 andx = 8, respectively. Altogether ten subgenera of the genusAster, five of them subdivided further into a total of 24 sections, are recognized as having representative species in the New World. All basionyms and type species are listed, and a number of new combinations and status changes are validated in accordance with the International Rules of Botanical Nomenclature. Where known (from literature and personal research), chromosome numbers are recorded for the species.     

The base number of ��loxoscaphoid�� Asplenium species and its implication for cytoevolution in Aspleniaceae

Background and Aims

‘Loxoscaphoid’ Asplenium species are morphologically a remarkably distinct group of Aspleniaceae. Except for two preliminary chromosome counts of Asplenium theciferum, the cytology of this group of species has, however, been largely unstudied.

Methods

Chromosome counts were obtained by acetocarmine squash preparations of one mitotic cell and several meiotic cells. Relative DNA content of gametophytic and sporophytic cells was determined by flow cytometry. The phylogenetic placement of A. loxoscaphoides, A. rutifolium s.l. and A. theciferum s.l. was investigated through an analysis of rbcL sequences.

Key Results

The dysploid base number is reported to be x = 35 in Asplenium centrafricanum, A. loxoscaphoides, A. sertularioides and A. theciferum. Analysis of rbcL sequences confirms that ‘loxoscaphoids’ nest robustly within Asplenium. Several high ploidy levels exceeding the tetraploid level were found in A. theciferum s.l. and A. rutifolium s.l. All taxa proved to be sexual.

Conclusions

Four base numbers are known at present for Aspleniaceae: x = 39, 38, 36 and 35. The dysploid base number x = 35 found in the ‘loxoscaphoid’ Asplenium spp. sheds a novel light on the cytoevolution of the whole family. We postulate a recurrent descending dysploid evolution within Aspleniaceae, leading to speciation at the (sub)generic and species/group level.     

A review of chromosome cytology in Hyacinthaceae subfamily Ornithogaloideae (Albuca, Dipcadi, Ornithogalum and Pseudogaltonia) in sub-Saharan Africa

The chromosome cytology of Hyacinthaceae subfamily Ornithogaloideae is reviewed within the framework of a recent molecular-based classification, with particular emphasis on its center of diversity in sub-Saharan Africa. We also provide new chromosome counts for sections that are unknown or poorly known cytologically. Albuca subgen. Namibiogalum (9 spp.) probably has an ancestral base number of x = 10 but subgen. Albuca (± 70 spp), subgen. Monarchos (9 spp.) and subgen. Osmyne (36 spp.) have x = 9. The pattern in subgen. Urophyllon (3 spp.) is remarkable: although x = 6 is likely, the species in the section exhibit a range of 2n = 12, 10, 8, 6 and 4 (exclusive of polyploidy). All karyotypes have three large chromosome pairs and a variable number of small chromosomes. Pseudogaltonia (2 spp.) has x = 9 and Dipcadi (26 spp.) possibly x = 9 in series Uropetalum and x = 6 in series Dipcadi, which exhibits a pattern of descending dysploidy leading to n = 3 in D. marlothii. In Ornithogalum (± 130 spp.) chromosome numbers are known for only 24 of the ± 84 sub-Saharan species, mostly from subgen. Aspasia and subgen. Ornithogalum sect. Linaspasia, both of which have x = 6, and from subgen. Galtonia, which has x = 8. In contrast, x = 7 is basic for the Eurasian sects. Honorius and Melophis, and x = 18 seems likely for sect. Cathissa. Sect. Ornithogalum, the cytology of which we does not examine in detail, may have x = 9. Polyploidy is apparently rare in the sub-Saharan African ornithogaloids, in marked contrast to the high frequency of polyploidy among Eurasian species. In Albuca just 3 or possibly 4 sub-Saharan species (9% or 13% of those counted) are exclusively polyploid and 5 more have diploid and polyploid races; and in sub-Saharan Ornithogalum, only the tropical O. gracillimum is exclusively polyploid, and the western southern African O. hispidum has diploid and polyploid races.     

Chromosome studies in southern species of Mimosa (Fabaceae,Mimosoideae) and their taxonomic and evolutionary inferences

In this work, chromosome numbers and karyotype parameters of 36 taxa of the genus Mimosa were studied, especially from the southern South America center of diversification. Results support that x = 13 is the basic chromosome number in the genus. Polyploidy is very frequent, ca. 56 % of the total of the studied species here are polyploid, confirming that polyploids are more frequent at higher latitudes. The most common ploidy levels found are 2x and 4x, but some species studied exhibit 6x and 8x. In different groups, several ploidy levels were found. Parameters of chromosome size show statistically significant differences between close species, and asymmetry index A ₂ exhibited low variation between them. It is possible to infer variations of chromosome size between diploids and tetraploids and between basal and derived taxa. The present studies confirm or reveal polyploidy in several groups of South America which are highly diversified in the southernmost area of distribution of the genus, such as sect. Batocaulon ser. Stipellares and sect. Calothamnos. Our data are discussed in a taxonomic context, making inferences about the origin of some polyploid taxa. Polyploidy could be an important phenomenon that increases the morphologic diversity and specific richness in southern South America. On basis of our data, it is possible to hypothesize hybridization between same-ploidy level or different ploidy level taxa. As already shown in the literature, our results confirm the importance of the polyploidy in the speciation of the genus.     

New species and combinations in Thelypteris subg. Goniopteris (Thelypteridaceae)

Three new species ofThelypteris are described and illustrated:T. littoralis andT. paranaensis, endemic to the rainforest of the southern Brazilian Atlantic coast, andT. multigemmifera, endemic to inland gallery forests of the São Paulo state. In addition, four new combinations are made:T. cutiataensis (Brade) Salino,T. iguapensis (C. Chr.) Salino. A new name is proposed forDryopteris lugubris var.quadrangularis:T. montana Salino. The species treated here belong toThelypteris subg.Goniopteris due to the indument of furcate and stellate trichomes.     

Chromosome numbers of Orthocarpus and related monotypic genera (Scrophulariaceae: Subtribe castillejinae)

A chromosome number ofn=12 is reported for the three monotypic genera of subtribe Castillejinae:Clevelandia beldingii, Gentrya racemosa, andOpicopephalus angustifolius. Chromosome numbers ofOrthocarpus correspond mostly with current infrageneric classification. SubgenusTriphysaria hasn=11.Orthocarpus sectionsCastillejoides andCordylanthoides, which are closely related toCastilleja (x=12) and the three monotypic genera above, haven=12 with aneuploid reductions ton=10 inO. linearilobus andn=11 inO. lacerus (a species also withn=12). Tetraploids are found in two species.O. brevistylus (n=24) andO. hispidus (n=12, 24). The polyploid.O. laciniatus (n=36, 48) of Peru is postulated to be of hybrid origin between a species ofCastilleja andOrthocarpus attenuatus. SubgenusOrthocarpus sectionOrthocarpus, which hasn=14 in all species except.O. bracteosus (n=15), stands apart both morphologically and in chromosome number from the remainder of the genus.     

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.     

Karyological study in the genusTuberaria sect.Scorpioides (Cistaceae): Taxonomic and evolutionary inferences

A polyploid complex ofTuberaria sect.Scorpioides species in S Spain has been studied. We have found the chromosome numbers n = 12 and 18, which probably originated from the extinct base number x = 6 by means of allopolyploidy. Octoploids (n = 24) seem to be the result of different crossings, and fall morphologically withinT. guttata, T. inconspicua, orT. commutata. Backcrosses between octoploids and tetraploids originate new hexaploids (n = 18), which, once stabilized, behave as amphidiploids. Most of these also fall within the traditional concept ofT. guttata, so that the enormous polymorphism of this taxon seems to be justified.     

Lobelia vivaldii (Campanulaceae: Lobelioideae), a remarkable new species of sect. Tylomium from Isla de Mona,Puerto rico

A new species ofLobelia (Campanulaceae: Lobelioideae) is described from Isla de Mona, a Caribbean island not previously known to harbor the genus. The small woody rosette-plants were collected in 1988 and 1992 at the northeastern corner of the island, where they grew from crevices in the vertical limestone cliffs, approximately 60 m above the sea. The new species,Lobelia vivaldii, belongs to sect.Tylomium, a group of approximately 36 suffruticose and woody species endemic to the Caribbean basin. Within that section, it is most similar to and apparently derived fromL. cirsiifolia, a less woody rain forest species found at higher elevations on several islands in the Lesser Antilles.     

Fertility relationships ofGeranium (Geraniaceae): sectt.Ruberta,Anemonifolia, Lucida andUnguiculata

Cross-pollinations were carried out among 11 briefly described species ofGeranium. Eight species pairs produced hybrids, of which five had not been reported before. The close relationship ofG. purpureum, G. robertianum andG. rubescens (sect.Ruberta) was confirmed; they form a polyploid series (diploid, tetraploid and octoploid on base x = 16). ForG. canariense (sect.Anemonifolia), another octoploid on base x = 16, the results suggest greater affinity with the former section.G. maderense andG. palmatum of sect.Anemonifolia (2n = 68) are confirmed as closely related to each other.G. maderense produced hybrids withG. robertianum (2n = 64; sect.Ruberta) and withG. cataractarum (2n = 36; sect.Unguiculata). Meiosis in the latter hybrid suggests allopolyploidy between parents with 2n = 32 and 2n = 36. Whereas all these species clearly form a very close alliance,G. lucidum (sect.Lucida) andG. macrorrhizum andG. dalmaticum (both sect.Unguiculata), appear genetically more isolated from them. One plant ofG. macrorrhizum ×G. dalmaticum was raised. — In crosses where hybrids or non-germinating seeds resulted, the reciprocal cross in the majority of cases produced a greatly inferior result or none at all. This asymmetry of response could in some cases be explained by inability of pollentubes from short-styled parents to reach the ovary of a long-styled species and in others by a modification of Hogenboom's theory of incongruity, but neither explanation works for every case. In all our asymmetric results where the ploidy level differed the diploid was the successful female, not the tetraploid, as is usually the case. — Some variation in results from year to year could be attributed to weather conditions.Dedicated to Hofrat Univ.-Prof. DrK. H. Rechinger on the occasion of his 80th birthday.     

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.     

The taxonomic significance of the fatty acid content of seeds of Linum

About sixty taxa in the genusLinum are arranged systematically and are compared using data published earlier for the fatty acid content of their seeds. Plants of sect.Linum, of sect.Dasylinum, and Old World species of sect.Linastrum generally have higher percentages of linolenic acid and lower percentages of linoleic acid; plants of sect.Syllinum, of sect.Cathartolinum, and New World species of sect.Linastrum have the reverse tendencies. The segregation ofL. stelleroides from other species of sect.Linum and the maintenance ofL. tenuifolium in a section separate from other tested species are supported.