Chloroplast DNA and isozyme evidence on the evolution ofSenecio vulgaris (Asteraceae)

Chloroplast DNA (cpDNA) and isozyme variation were analyzed over a range of populations of two infraspecific taxa of the tetraploidSenecio vulgaris. The isozyme data were supportive of the hypothesis that the weedy and cosmopolitanS. vulgaris var.vulgaris is an evolutionary derivative ofS. vulgaris subsp.denticulatus from the coasts of W Europe and montane altitudes in S Spain and Sicily. The two taxa exhibited a very high genetic identity with subsp.denticulatus containing slightly more isozyme diversity than was found in var.vulgaris. — Three cpDNA haplotypes (A, B, C) already known from other Mediterranean diploid species ofSenecio were resolved in var.vulgaris, and an additional fourth haplotype (E) was found in subsp.denticulatus. Two alternative hypotheses were chosen to account for the origin and maintenance of the observed cpDNA composition ofS. vulgaris. It either reflects (1) the retention of an ancestral polymorphism which stems from the recurrent and polytopic formation of ancestral tetraploid lineages; or (2)S. vulgaris originally was characterized by haplotype E, and haplotypes A, B and C were acquired through repeated introgressive hybridization with related diploid species. The finding that very low levels of nuclear (isozyme) diversity were present in both taxa ofS. vulgaris examined supports the second of these two hypotheses; however, more detailed analysis of nuclear genetic diversity is required before a firm conclusion can be reached on this matter.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Isozyme differentiation in the endemic genusArgyranthemum (Asteraceae: Anthemideae) in the Macaronesian Islands

The genusArgyranthemum (Asteraceae: Anthemideae) comprises 38 taxa and is restricted to the archipelagos of the Canaries, Selvagens, Madeira, and Desertas in the Macaronesian biogeographic region. An electrophoretic study, including 17 enzyme loci and at least one population of each of the described taxa, was carried out. High identity (low distance) values between taxa (mean of 0.893) were obtained despite the old age of the islands, their close proximity to the African continent, and the fact thatArgyranthemum is the most species-rich and variable genus in Macaronesia. These results suggest that the genus is monophyletic and that it has evolved very rapidly in these islands. There is little correspondence between taxonomy and neighbor-joining analysis based on Roger's genetic distances, but in several instances populations from the same islands cluster together despite being from different species or even different sections. It is suggested that repeated genetic bottlenecks associated with the founding of new populations during radiation of the genus resulted in lineage sorting of ancestral allozyme polymorphisms. Because every population has a high average identity with all other populations, lineage sorting could result in populations of different taxa being slightly more similar than populations of the same taxon. Gene flow between different species on the same island could account for some populations clustering by island of origin rather than taxonomic disposition. Average allozyme diversity within populations (0.098) is 50% higher than the mean total diversity for species endemic to oceanic islands.     

Molecular phylogeny ofCheirolophus (Asteraceae:Cardueae-Centaureinae) based on ITS sequences of nuclear ribosomal DNA

The genusCheirolophus has an interesting western Mediterranean and Macaronesian distribution. Here we investigate the delimitation of the genus and its exclusion from the large genusCentaurea, the systematic position of the related genusPaleocyanus, the delimitation of some species and the phylogeny of the group. We have carried out a phylogenetic analysis of the PCR-generated sequences of the internal transcribed spacers (ITS-1 and ITS-2) of the nuclear ribosomal DNA. The results suggest that the genus, includingPaleocyanus crassifolius is monophyletic; thus, a new combination of this species underCheirolophus is proposed. The Macaronesian group of species is also monophyletic, indicating a single colonization of the archipelago. The poor resolution of microspecies in the Macaronesian group reinforces the hypothesis of a very recent differentiation of the group.     

Adaptive radiation and genetic differentiation in the woodySonchus alliance (Asteraceae:Sonchinae) in the Canary Islands

The woodySonchus alliance consists of 19 species ofSonchus subg.Dendrosonchus, one species ofSonchus subg.Sonchus and species of five genera (i.e.Babcockia, Sventenia, Taeckholmia, Lactucosonchus, Prenanthes), and is restricted primarily to the archipelago of the Canaries in the Macaronesian phytogeographical region. An enzyme electrophoretic study, including 13 loci, was conducted to assess genetic diversity within and divergence among species of the alliance. Nei's genetic identities (distances) between genera and/or subgenera range from 0.490 (0.714) to 0.980 (0.013), and pairwise comparisons of all populations show relatively high genetic identities, with a mean of 0.804. The high identities further support the genetic cohesiveness of the alliance and its single origin on the Macaronesian islands. Species in the alliance also show about 50%; higher total genetic diversity (H_T) than the mean for other oceanic endemics. There is greater divergence between endemics or species on older islands compared to those on younger islands, which suggests that time is a factor for divergence at allozyme loci. Furthermore, populations on older islands have higher total genetic diversities and lower identities than conspecific populations on younger islands. These results imply early colonization, radiation, and divergence of the woodySonchus alliance on older islands followed by subsequent colonization to younger islands. The taxonomic distribution of alleles in the alliance indicates lineage sorting also played a role in divergence among species. Lineage sorting may also produce nonconcordance with either taxonomic designation or the pattern of variation obtained from other molecular markers such as ITS sequences of nrDNA. Timing for the origin and radiation of the alliance agrees with the estimate based on ITS sequences, and suggests that the early divergence and rapid radiation took place during the Late Tertiary on either Gran Canaria or Tenerife.     

Isozyme evidence and the origin ofSenecio vulgaris (Compositae)

An electrophoretic survey of isozyme variation was conducted to test the hypothesis thatSenecio vulgaris L. (2n = 40) is of autotetraploid origin fromS. vernalis Waldst. & Kit. (2n = 20). It was established thatS. vulgaris exhibited fixed heterozygosity at three loci examined, showed disomic inheritance at all polymorphic loci, and contained a gene (Est-1) and an allele (Aat-3b) which were not present in the single population ofS. vernalis surveyed. From this it is concluded thatS. vulgaris is not of autotetraploid origin. Instead, the genetic evidence is in keeping with an allopolyploid origin ofS. vulgaris with the possibility thatS. vernalis acted as one of its two parents.     

The origin ofSenecio vulgaris (Asteraceae)

Senecio vulgaris L. (2n = 40) is suggested to be of autotetraploid origin fromS. vernalis Waldst. & Kit. (2n = 20). This conclusion is based on results obtained from experimental hybridisations and cytological observations, and the consideration of morphological affinities, patterns of geographical distribution and hybrid formation under natural conditions. The morphological differences between the two species are related to a difference in the breeding system. WhilstS. vernalis is self-incompatible,S. vulgaris is self-compatible and strongly self-pollinating. Equally, other self-pollinating taxa traditionally associated withS. vulgaris are shown to have evolved independently from outbreeding relatives. Within a narrow frame of relationship, annual weeds have evolved in different cytotaxonomic circumstances.     

A cladistic study of North AmericanCoreopsis (Asteraceae: Heliantheae)

A cladistic study of all 44 species of North AmericanCoreopsis was performed using 35 characters. The resulting cladogram indicated that all 11 sections are monophyletic. At the intersectional level, two lineages were revealed, one consisting of six sections occurring almost exclusively in Mexico and California, and another comprising five sections restricted largely to the eastern and southeastern United States. The cladogram is similar to phylogenies produced by less explicit methods but it differs in two major respects: the monotypic sect.Silphidium is placed with other sections from the southeastern United States rather than with Mexican sections, and sect.Anathysana from Mexico is more closely allied with the three California sections than with sect.Electra from Mexico.     

Intraspecific evolution ofMicroseris pygmaea (Asteraceae,Lactuceae) analyzed by cosegregation of phenotypic characters (QTLs) and molecular markers (RAPDs)

The Chilean annual,Microseris pygmaea, has differentiated in distinct coastal and inland series of populations after long-distance dispersal from western North America. Two plants from the most diverse biotypes were crossed, a large F₂ was raised and analysed for segregation of 30 phenotypic characters. Segregation of molecular markers (47 RAPDs, 1 RFLP, 2 isozymes) was determined in a subpopulation of 45 plants which include all extremes for the phenotypic characters. 32 marker/character cosegregations were significant at the 1% level in t-tests between dominant and homozygous recessive marker genotypes. Considering linkage among markers and pleiotropy of certain marker loci, the number of independent quantitative trait loci (QTLs) is reduced to about 18. Interactions among 2 or 3 QTLs affecting one character have been characterized. The phenotypic differentiation ofM. pygmaea during its evolution from a single founder individual begins to be understood at the level of single-gene mutants.     

The genusPicris (Asteraceae,Lactuceae) in Tropical Africa

Three species of the genusPicris L. are native in Tropical Africa:P. abyssinica Sch. Bip. (Ethiopia),P. xylopoda Lack, spec. nova (Nigeria, Ethiopia) andP. humilis DC. (Senegal, Mali). There are indications that the two perennial species,P. abyssinica andP. xylopoda, are related to and have evolved from a primitive Central Asiatic stock in a manner parallel to many African species ofCrepis L.P. humilis, on the other hand, is a small annual plant with a high number of derived characters. The introduced species of European origin growing south of the Sahara are briefly mentioned.     

Floral rewards in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The genusScrophularia in the Iberian Peninsula and Balearic Islands comprises two sections,Scrophularia andCaninaG. Don. Analyses were carried out on flower production, flower duration and their sexual phases, pollen and nectar production together with observations on their pollinators. Nectar production is correlated with corolla size and pollen production with anther size. The taxa of sect.Scrophularia show greater nectar and pollen production than those of sect.Canina. Also, those of the first section produce more ovules per ovary than those of the second group, production being correlated with the ovary size.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and the Balearic Islands. I.     

Taxonomy and phylogeny of the tribePlucheeae (Asteraceae)

The tribePlucheeae (Benth.)A. Anderb., has been analysed cladistically by means of a computerized parsimony program (Hennig 86), using theArctotideae as outgroup. The results of the analysis are presented in a consensus tree and one cladogram. Four major monophyletic subgroups can be recognized: TheColeocoma group (3 genera), thePterocaulon group (3 genera), theLaggera group (6 genera), and thePluchea group (12 genera). All recognized genera are described and most genera are supplied with taxonomical notes including comments on their taxonomic status. Genera such asBlumea, Pluchea, andEpaltes are demonstrated to be unnatural assemblages.Monarrhenus andTessaria are both closely related to thePluchea complex. The old generic nameLitogyne Harv. has been taken up for one species ofEpaltes, the genusRhodogeron is reduced to a synonym ofSachsia, and the following new combinations are made;Litogyne gariepina (DC.)A. Anderb., andSachsia coronopifolia (Griseb.)A. Anderb.     

Taxonomy ofCarminatia (Asteraceae,Eupatorieae)

A taxonomic study of the largely Mexican genusCarminatia is rendered. It is comprised of three closely related species:C. tenuiflora, C. recondita andC. anomala spec. nova. Illustrations, dot maps, keys to species and complete synonymy are presented.     

Biology of afroalpineDendrosenecio (Asteraceae)

The genusDendrosenecio (giant groundsels), encompassing three species and 12 subspecies, is endemic to the high mountains of East and Central Africa where it constitutes the most conspicuous components of the afroalpine vegetation. Two lifeforms, the arborescent and the prostrate rhizomatous, are regarded as the results of evolution from forest-living woody or herbaceous ancestors. Due to the uninterrupted growth period in the tropics, there are no anatomical or morphological features which allow conventional age determination. However, stem elongation rates have been determined (3–5.5 cm per year) and indicate an age of about 250 years for the tallest arborescent Dendrosenecios which may reach a height of 10 m. 30 to 120 large leaves are clustered in an enormous terminal rosette, justifying the term giant rosette plants. A leaf bud, consisting of about as many developing leaves as the rosette contains, is found in the center. During the nocturnal frost period the adult rosette leaves form a so-called night-bud by nyctinastic upwards bending and thus protect the leaf bud from freezing by insulation. The stem is surrounded by a mantle of persistent dead leaves; this ameliorates the microclimate of the pith-cells which greatly contribute to water transport into the leaves. Below the leaf rosette a zone of putrefaction is found, from where the decay of the dead leaves apparently provides nutrients directly to the growing stem. The population dynamics of the arborescentD. keniodendron is characterized by a simultaneous inflorescence development at irregular intervals of up to more than twenty years. Due to sporadic flowering and a seedling survival rate of less than 1%, oscillations of the population size are to be expected.     

Self-fertility and associated flower head traits in the Iberian taxa ofLactuca and related genera (Asteraceae: Lactuceae)

Sexual reproduction is the main reproduction mechanism among the 14 wild Iberian taxa ofLactuca, Prenanthes, Cicerbita, andMycelis. High levels of self-fertilization occur inLactuca, as well as facultative and obligate xenogamy. Xenogamy is strongly correlated with large capitula having blue or bright yellow colouration, high P/O ratios, and long anthesis, whereas self-fertilization is correlated with smaller capitula having pale yellow colouration, small P/O ratios, and short anthesis. Large variations occur in P/O index among taxa showing similar fertilization mechanism, probably in relation to the number of florets included in flower heads. Interesting differences in reproductive systems have been detected between subspecies ofLactuca viminea.     

Comparison of chloroplast and nuclear phylogeny in the autogamous annualMicroseris douglasii (Asteraceae: Lactuceae)

Morphology suggests that the Californian annualMicroseris douglasii is a monophyletic sister group to the other three diploid annuals ofMicroseris. Phylogenetic analysis of 44 inbred strains ofM. douglasii derived from 23 populations with 72 RAPD markers in the nuclear DNA strongly supports this phylogeny. However, 13 chloroplast RFLPs divideM. douglasii into four distinct groups. Two of these each share one or more cpRFLPs withM. bigelovii andM. pygmaea. Several hypotheses can explain the incongruence between nuclear and chloroplast phylogeny: (1) random sorting out of chloroplasts during phylogeny from a polymorphic pool, (2) cytoplasmic introgression from the related annualM. bigelovii intoM. douglasii after hybridization followed by elimination of theM. bigelovii nuclear genome. We suggest cytoplasmic introgression as the most likely origin. Possible remnants of nuclear introgression have been found in two populations ofM. douglasii that are polymorphic for chloroplast types. In these populationsM. bigelovii type chloroplast DNA seems to be accompanied by nuclear genes for flower color and leaf shape.     

Karyosystematic studies on threeCrepis species (Asteraceae) endemic to Greece

This work examines the cytogeographical distribution, the morphological characters, and the karyotypes of threeCrepis species endemic to Greece (C. sibthorpiana, C. incana, andC. heldreichiana). C. sibthorpiana is diploid (2n = 2x = 8),C. incana is diploid (2n = 2x = 8) and tetraploid (2n = 4x = 16, 17), andC. heldreichiana is always dekaploid (2n = 10x = 40). The Giemsa positive bands, usually pairs of dots, are mainly centromeric inC. incana, while they are terminal inC. sibthorpiana (on the short arm of all chromosomes) and inC. heldreichiana (on both arms of all chromosomes). Intercalary C-bands are scarce and usually variable within karyotypes, individuals, and species. The most variable karyotype both in Feulgen and Giemsa preparations is that ofC. incana, in which also supernumerary chromosomes were observed, which are polysomic to standard set members. On the basis of morphological and karyological data the evolutionary relationships among the threeCrepis taxa are discussed.     

Genetic analysis and conservation of the endangered Canary Island woody sow-thistle, Sonchus gandogeri (Asteraceae)

Sonchus gandogeri, a woody sow-thistle, is an endangered Canary Island endemic with only two known populations, one in the El Golfo and another in the Las Esperillas of El Hierro. Amplified fragment length polymorphism (AFLP) markers were used to assess the genetic variation within and among populations. The mean genetic diversity of two populations was estimated to be 0.380, and the El Golfo population (0.380) had higher genetic diversity than the southeastern one (0.268). The unbiased Neis genetic identity between the two populations was 0.846. The mean genetic diversity of S. gandogeri was much higher than that of the other endangered plant species. This is perhaps due to breeding system, life form, extinction, and/or introgressive hybridization and hybrid origin of the taxon. This study also indicates that the two populations are not strongly differentiated (G_ST=0.149). This study suggests that S. gandogeri is more likely to become extinct due to environmental or demographic forces than genetic factors, such as inbreeding depression. More strict control of introduced herbivores is necessary to protect these populations, and germplasm collection for ex situ conservation is needed.     

Genotype relationships inMicroseris elegans (Asteraceae,Lactuceae) revealed by DNA amplification from arbitrary primers (RAPDs)

The genetic relationships among 10 inbred lines representing 10 populations of the autogamous annualMicroseris elegans from throughout California has been determined using random amplified polymorphic DNAs (RAPDs). Seventeen arbitrary 10 base pair primers produced 134 amplification products; 81 of these were shared by two or more strains. The 3 genotypes from Northern California are closely related as are 3 genotypes from Middle Californian populations which are not nearest neighbors. DNA fingerprinting with the oligonucleotide (GATA)₄ gave compatible results, but the comparison was limited to samples run on one gel. Isoenzyme patterns are compatible with the DNA results, but limited by the very low number of informative polymorphisms. The clustered relationship among genotypes within a species and their geographic distribution suggests very restricted genetic recombination and an origin of new populations from randomly dispersed achenes within the range of the species.