A phylogenetic study ofSaxifraga sect.Saxifraga (Saxifragaceae) based on nrDNA ITS sequences

Parsimony analyses of 54 nrDNA ITS (Internal Transcribed Spacer) sequences ofSaxifraga sect.Saxifraga were performed. In addition to some unresolved clades, there is strong disagreement between the ITS phylogeny and previous classifications based primarily on morphology. The extensive cytological instability of sect.Saxifraga prevents previous cytotaxonomical results from resolving the incongruence between molecular and morphological data. Dissimilar topologies between chloroplast (matK) and nuclear (ITS) trees for eight species of sect.Saxifraga suggest that gene trees and the true species tree are not coincident. Recent and mid-term reticulation is proposed as an explanation for the incongruence between morphological, cytological, organellar, and nuclear data. Homogenization in multigene families, such as the ITS region, via concerted evolution may be the key to the interpretation of results based on ITS sequences within sect.Saxifraga. The use of organellar genes in a larger sample should help to determine whether extensive reticulation occurs in sect.Saxifraga, as has been documented in various genera of Saxifragaceae.     

Phylogenetic relationships among Lactuca (Asteraceae) species and related genera based on ITS-1 DNA sequences

Internal transcribed spacer (ITS-1) sequences from 97 accessions representing 23 species of Lactuca and related genera were determined and used to evaluate species relationships of Lactuca sensu lato (s.l.). The ITS-1 phylogenies, calculated using PAUP and PHYLIP, correspond better to the classification of Feráková than to other classifications evaluated, although the inclusion of sect. Lactuca subsect. Cyanicae is not supported. Therefore, exclusion of subsect. Cyanicae from Lactuca sensu Feráková is proposed. The amended genus contains the entire gene pool (sensu Harlan and De Wet) of cultivated lettuce (Lactuca sativa). The position of the species in the amended classification corresponds to their position in the lettuce gene pool. In the ITS-1 phylogenies, a clade with L. sativa, L. serriola, L. dregeana, L. altaica, and L. aculeata represents the primary gene pool. L. virosa and L. saligna, branching off closest to this clade, encompass the secondary gene pool. L. virosa is possibly of hybrid origin. The primary and secondary gene pool species are classified in sect. Lactuca subsect. Lactuca. The species L. quercina, L. viminea, L. sibirica, and L. tatarica, branching off next, represent the tertiary gene pool. They are classified in Lactuca sect. Lactucopsis, sect. Phaenixopus, and sect. Mulgedium, respectively. L. perennis and L. tenerrima, classified in sect. Lactuca subsect. Cyanicae, form clades with species from related genera and are not part of the lettuce gene pool.     

Frequent gain and loss of pyrrolizidine alkaloids in the evolution of Senecio section Jacobaea (Asteraceae)

Pyrrolizidine alkaloids (PAs) of the macrocyclic senecionine type are secondary metabolites characteristic for most species of the genus Senecio (Asteraceae). These PAs are deterrent and toxic to most vertebrates and insects and provide plants with a chemical defense against herbivores. We studied the PA composition of 24 out of 26 species of Senecio section Jacobaea using GC-MS. The PA profiles of eight of these species have not been studied before and additional PAs were identified for most other species that were included in previous studies. With one exception (senecivernine) all 26 PAs identified in sect. Jacobaea can be regarded as derivatives of the biosynthetic backbone structure senecionine. Based on the PA profiles of the species of sect. Jacobaea and the results of previous tracer studies, we constructed two hypothetical biosynthetic scenarios of senecionine diversification. Both scenarios contain two major reactions: the conversion of the necine base moiety retronecine into the otonecine moiety and site-specific epoxidations within the necic acid moiety. Further reactions are site-specific hydroxylations, sometimes followed by O-acetylations, site-specific dehydrogenations, E, Z-isomerizations, and epoxide hydrolysis and chlorolysis. The GC-MS data and both biosynthetic scenarios were subsequently used to study the evolution of PA formation in sect. Jacobaea by reconstructing the evolutionary history of qualitative PA variation in this section. This was achieved by optimizing additive presence/absence data of PAs and types of enzymatic conversions on a maximum parsimony cladogram of section Jacobaea inferred from DNA sequence and morphological data. Besides showing large intra- and interspecific variation, PA distribution appears to be largely incidental within the whole clade. These results together with the finding that all but one of the PAs identified in sect. Jacobaea are also present in species of other sections of Senecio indicate that differences in PA profiles in Senecio can not be explained by the gain and loss of PA specific genes, but rather by a transient switch-off and switch-on of the expression of genes encoding PA pathway-specific enzymes.     

Three new species of Senecio (Asteraceae) from Mexico

Senecio moranii, Senecio martirensis, andSenecio semperamatae, all from Mexico, are described as new.     

Phylogenetic position of Mediterranean Astereae and character evolution of daisies (Bellis,Asteraceae) inferred from nrDNA ITS sequences

Phylogenetic analyses of nrITS sequences of Asteraceae revealed that the Bellis group is a natural assemblage comprising all the species of Bellis and Bellium, but not Rhynchospermum. In contrast, we propose to include the genera Bellis, Bellium, and Bellidastrum in the subtribe Bellidinae in the interest of circumscribing natural groups. Our results also suggest an early diversification in the western Mediterranean Basin of two monophyletic lineages, Bellis and Bellium. Three major groups can be distinguished within BELLIS: (1) the B. perennis group, containing five annual and perennial species with three ploidy levels (diploid, octoploid, and decaploid), which are distributed throughout the Mediterranean Basin despite lack of pappus; (2) the Bellis sylvestris group, with five annual and perennial species primarily from the western Mediterranean, in which there are five ploidy levels (diploid, tetraploid, hexaploid, octoploid, and decaploid); and (3) a basal grade consisting of three diploid, perennial species which displays remarkable diversification of morphologies. Striking characteristics, such as an annual life form, polyploidy, and loss of pappus, seem to have occurred in parallel several times and in different geographical areas during the early diversification of Bellis species in the western Mediterranean. Character evolution reconstructions allow us to describe a putative ancestor of the genus Bellis (proto-Bellis).     

A new species of Anthemis sect. Hiorthia (Asteraceae) from SE Sicily

A new species of Anthemis sect. Hiorthia (Asteraceae) is described from Sicily. This taxon, named A. piagnattiorum, is confined to a restricted area of the Hyblaean territory (SE Sicily), in the Natural Reserve of Cava Grande (Cassibile, Siracuse), where it takes shelter on carbonate rocky habitat and participates, with other Sicilian endemic taxa, to the composition of a termophilous chasmophytic community (Putorio calabricae–Micromerietum microphyllae).     

A phylogenetic study ofPolygonum sect.Tovara (Polygonaceae) based on ITS sequences of nuclear ribosomal DNA

Polygonum sect.Tovara comprises three morphologically very similar species;P. virginianum,P. filiforme, andP. neofiliforme. Sequences of internal transcribed spacers (ITSs) of nuclear ribosomal DNA of these were determined to examine phylogenetic relationships and the levels of differentiation among them. The size of ITS 1 was 241 bp inP. filiforme andP. neofiliforme, and 242 bp inP. virginianum. The size of ITS 2 was 243 bp, and that of the 5.8S rRNA coding region was 163 bp. The ITS sequences clearly separate North AmericanP. virginianum from the eastern Asian species. Nucleotide divergence between them ranges from 3.3% to 3.8% for ITS 1 and from 9.3% to 10.7% for ITS 2. The molecular data also revealed that two eastern Asian species are closely related but should be treated as distinct species.     

Numerical morphological analysis of Lettuce cultivars and species (Lactuca sect.Lactuca,Asteraceae)

L. sativa L. cultivars were compared morphologically with accessions ofL. saligna L.,L. serriola L., andL. virosa L. in order to define the infra- and interspecific variation. Multivariate analysis (principal component analysis) distinguished four groups corresponding with the four species. Infraspecific variation (e.g., colour, leaf shape) withinL. sativa is the largest, because breeding has caused a large variation in vegetative features; infraspecific variation withinL. serriola is also large. Three plants of one accession were placed betweenL. sativa andL. serriola because they showed morphological characteristics of both species. They are still classified asL. sativa. The morphological differences betweenL. sativa andL. serriola are too large to consider these two as one species. The descent of the cultivar groups and the four species is described.     

Two new species of Senecio section Palmatinervii (Asteraceae) from eastern Mexico

Two closely related new species,Senecio grimesii andS. marquezii, belonging to the sectionPalmatinervii are described, illustrated, and compared to their nearest relative,S. heterogamus. An erratum to this article is available at .     

Molecular phylogenetic position of Japanese Abies (Pinaceae) based on chloroplast DNA sequences

The phylogenetic positions of Japanese Abies species (A. firma, A. homolepis, A. veitchii, A. sachalinensis, and A. mariesii) were revealed based on nucleotide sequences of chloroplast DNA. First, rbcL sequences of 24 Abies species worldwide were analyzed to clarify the phylogenetic position of the Japanese Abies within this genus. While 4 of the Japanese species formed a monophyletic group with almost no sequence divergence, A. mariesii was placed into a clearly different group with North American species. Second, to clarify the relationships among the species in the monophyletic group, sequences of rbcL, matK, and six spacer regions (total ca. 5 kb) were analyzed for 12 species of the group, with A. mariesii as an outgroup. Although a unique sequence and some species-specific sequences were detected in A. mariesii and in A. firma and A. homolepis, respectively, A. veitchii and A. sachalinensis gave identical sequences for all the sequenced regions.     

Recurrent origin of peripheral,coastal (sub)species in Mediterranean Senecio (Asteraceae)

Background: It is argued that coastal endemic taxa may evolve in parallel at the periphery of the distributional range of a widespread species.

Aims: We tested this hypothesis for the origins of three peripheral, coastal isolates of Senecio, S. glaucus ssp. glaucus (Israel), S. g. ssp. coronopifolius p.p. (Sicily), and S. hesperidium (Morocco), from widespread S. glaucus ssp. coronopifolius. We also determined the relative roles of selection vs. genetic drift in shaping phenotypic divergence in ssp. glaucus and S. hesperidium, using Lande’s test of neutral morphological change.

Methods: We surveyed morphological and/or allozyme variation in the three peripheral isolates and mainly inland populations of S. g. ssp. coronopifolius.

Results: Genetic data supported independent origins of the coastal taxa from nearby populations of ssp. coronopifolius. These descendant and ancestral populations showed pronounced morphological but weak genetic differentiation. Phenotypic similarities between ssp. glaucus (Israel) and S. hesperidium (Morocco) in plant height and floral traits may have resulted from parallel divergent selection from ssp. coronopifolius, though drift remains an alternative cause in S. hesperidium.

Conclusions: Our results indicate parallel ecotype formation and (sub)speciation in Senecio in which primarily selective vs. neutral determinants promoted the recurrent origin of coastal types in, respectively, Israel and Morocco.     

Molecular phylogenetic position of the genera Stephanonympha and Caduceia (Parabasalia) inferred from nuclear small subunit rRNA gene sequences

Nuclear small subunit (SSU) rRNA gene sequences were obtained by polymerase chain reaction from trichomonad symbionts of termites that belong to the Devescovinidae (Caduceia versatilis) and polymastigont Calonymphidae (Stephanonympha nelumbium). The unidentified SSU rRNA sequence Nk3, previously obtained from the termite Neotermes koshunensis, has also been shown to derive from a Stephanonympha sp. by in situ hybridization. These sequences were analysed in a broad phylogeny including nearly all identified parabasalid sequences available in the databases, and some as yet unidentified sequences likely deriving from the new order Cristamonadida (Devescovinidae, Calonymphidae, and hypermastigids Lophomonadida). A global phylogeny of parabasalids reveals a partial agreement between the clades identified in this work and the last classification of this phylum into four orders. However, this classification is still incongruent with our data and new taxonomic considerations are proposed. The analysis confirms the monophyly of the Cristamonadida and separates this order into two groups: the first unites nearly all the Devescovinidae including Caduceia and the Calonymphidae Coronympha and Metacoronympha, whereas the second group is composed of a few Devescovinidae, Lophomonadida, and Calonymphidae such as Stephanonympha. Caduceia is closely related to Devescovina, corroborating the marked morphological similarity between these two genera whereas Stephanonympha groups together with the Calonymphidae Snyderella and Calonympha. These data also confirm the polyphyly of the families Devescovinidae and Calonymphidae and support the arrangement of the axostyle-pelta complexes as a valuable character for taxonomic considerations within the Calonymphidae.     

Tortula and some related genera (Pottiaceae, Musci): phylogenetic relationships based on chloroplast rps4 sequences

 According to chloroplast rps4 sequence data the genus Syntrichia forms a monophyletic clade clearly separated from Tortula, while Pottia is shown to be polyphyletic and the Rhynchostegiae species (with rostrate lid) are very similar to Tortula. Crossidium is probably polyphyletic. The close affinity of Desmatodon, Stegonia, Pterygoneurum and some of the species of Phascum with Tortula is confirmed by the molecular data. Received May 23, 2002; accepted August 23, 2002 Published online: November 22, 2002 Address of the authors: O. Werner, R. M. Ros, M. J. Cano, J. Guerra (E-mail: werner@um.es) Departamento de Biología Vegetal (Botánica), Facultad de Biología, Universidad de Murcia, Campus de Espinardo, E-30100-Murcia, Spain.     

Hieracium busambarense,a new species of the sect. Grovesiana (Asteraceae) from Sicily (Italy)

Hieracium busambarense (Asteraceae), a new species from calcareous-dolomite cliffs of Rocca Busambra (western Sicily, Italy) is described and illustrated here.     

Chromosome numbers in selected species ofHieracium sect.Alpina (Asteraceae) from Central and eastern Europe

Chromosome numbers are reported for 15 taxa ofHieracium sect.Alpina (Griseb.)Gremli from Central and eastern Europe (Austria, the Czech Republic, Poland, Romania, Slovakia, Ukraine). The mode of reproduction was also studied for some of the taxa. For the first time the chromosome counts of 5 taxa from theHieracium rohacsense group are given:H. rohacsense Kit. (the West Carpathians),H. ratezaticum (Nyár. etZahn)Mráz (the South Carpathians) and a still unnamed taxon of theH. rohacsense group from Mt. Pop Ivan (the East Carpathians) are tetraploid (2n=36);H. rauzense Murr (the Eastern Alps) andH. borsanum Mráz (the East Carpathians) are triploid (2n=27).H. krivanense (Woł. etZahn)Schljakov (the West Carpathians) is tetraploid,H. brevipiliferum Mráz (the South Carpathians) is triploid; these are the first karyological reports of both taxa belonging to theH. fritzei group. The tetraploid chromosome number was revealed for the first time in an unnamed taxon of theH. nigrescens group from the West Carpathians, and inH. nigrescens subsp.koprovanum Rech. f. etZahn. InH. alpinum L. s.str. triploid (2n=27) populations from the West Carpathians and diploid populations (2n=18) from the East Carpathians were confirmed. ForH. halleri Vill. (theH. alpinum group, the West Carpathians) andH. nigrescens Willd. (the West Sudeten), the numbers 2n=27 and 2n=36, respectively were found, which is in accordance with previous data. The triploid level (2n=27) forH. pinetophilum (theH. fritzei group) and the tetraploid level (2n=36) forH. stygium Uechtr. (theH. chlorocephalum group) both from the West Carpathians were confirmed. One new species, onenomen novum and one new combination at the level of species are published in this paper.     

Systematics ofNama (Hydrophyllaceae): Flavonoids and phyletic position of sect.Arachnoidea and sect.Cinerascentia

Eight flavonoids, four 6-oxygenated flavones, two methyl ethers of luteolin, apigenin 6,8-C-diglucoside and quercetin 3-O-glucoside, were isolated fromNama lobbii andN. rothrockii, sole members of sects.Arachnoidea andCinerascentia, respectively. Both taxa diverge markedly from other namas in morphology and chromosome number and their placement inNama has been questioned. The occurrence of 6-oxygenated flavones in these taxa adds to their already distinctive nature. Flavonoid evidence argues that both are more closely allied toEriodictyon than either is toNama.     

Chloroplast DNA variations in sect. Cyclachaena of Iva (Asteraceae)

Much argument has been occasioned by questions having to do with generic concepts in the tribe Heliantheae, subtribe Ambrosiinae, especially involving the genus Iva. Some workers recognize five genera (e.g., Leuciva, Chorisiva, Oxytenia, Cyclachaena, and Iva s. str.), while others recognize only a single genus Iva s. 1. Although chemical, pollen ultrastructure, and other morphological data have challenged the latter concept, relationships among the segregate taxa are still unclear. The current study used cladistic analyses of chloroplast DNA variation to investigate evolutionary relationships within Iva s. l., especially within the controversial section Cyclachaena. Nuclear rDNA was also utilized to test for hybrid origin of selected taxa. The results support the dismemberment of Iva s. l. and recognition of Iva s. str., largely because Iva s. l. is shown to be paraphyletic. Most members of the section Cyclachaena were found to have relatively close interspecific relationships. Nevertheless, cpDNA data strongly support two lineages within Cyclachaena. One lineage, including I. xanthifolia, I. acerosa, I. nevadensis, and I. dealbata, has a close relationship with the genus Euphrosyne. The second lineage includes only one species, I. ambrosiaefolia, which is closely related to Dicoria. The origin of I. ambrosiaefolia was shown to be monophyletic, not involving hybridization.